Skip to main content

Full text of "Memoirs of the National Museum of Victoria"

See other formats


Memoirs Of the Number 37 

NATIONAL MUSEUM 

Of Victoria Melbourne Australia 30 June 1976 
V 



V 




f /i 



'JZsSei 



- i m^i 



r#1 



:M 



■t&&2J'-~:--;<:M 



■9 — 



3 



■ . 
■ 



■*s 





.•* 



&&&& -v 



» SJ 



Cover — An Obulkarra woman being painted with a clan 
design after death. 



MEMOIRS 

of the 

NATIONAL MUSEUM OF VICTORIA 

MELBOURNE AUSTRALIA 



No. 37 



Director 
J. McNally 

Deputy Director 
Thomas A. Darragh 

Editor 
Douglas M. Stone 



PUBLISHED BY ORDER OF THE TRUSTEES 
30 JUNE 1976 



Brown Prior Anderson Pty Ltd 5 Evans Street Burwood 3125 



NATIONAL MUSEUM OF VICTORIA 

Members of Council 

Sir Robert Blackwood, MCE BEE FIE Aust (President) 
Henry G. A. Osborne, BAgrSc (Deputy President) 
James C. F. Wharton, BSc (Treasurer) 

Professor E. S. Hills, CBE PhD (Lond) Hon DSc (Dunelm) DSc FIC FAA 

FRS 
Professor Sir Sydney Sunderland, CMG MD BS DSc FRACP FRACS FAA 
Dr. H. K. Worner, DSc (Melb) Hon DSc (Newcastle) ABSM FAA FRACI 

FIM F1MM MA1MM MAIME FTS 
Sir Henry Somerset, CBE MSc FRACI MAIMM 
Brigadier P. P. Jackson, CMG CBE BE FIE (Aust) FI Mech E FAIM 
Professor J. W. Warren, MA PhD (Calif) 

John McNally, ED MSc, Chief Executive Officer 
W. L. Drew, Secretary to Council 

Staff (as at 23/2/1976) 

Director: John McNally, ED MSc AMAA 

Deputy Director: Thomas A. Darragh, MSc DipEd AMAA 

Administration: 

A. G. Parsons (in charge) 

E. J. Peat 

E. Rowley 

Patricia Batchelor 

Enid Howell 

Kayelene Voege 

Lynette Anderson 

Scientific Staff 

Geology and Palaeontology: 

Curator of Fossils: T. H. Rich, PhD BA 

Curator of Minerals: W. D. Birch, BSc (Hons) 

Assistant Curator of Fossils: Vacant 

Assistants: F. Bussat, I. Stewart, FRMIT (Geol), Susan Gibson 

Vertebrate Zoology: 

Curator of Vertebrates: Joan M. Dixon, BSc (Hons) 
Curator of Birds: A. R. McEvey, BA 
Assistants: Linda Chegwidden, BSc, I. T. Beck 

Herpetology: A. J. Coventry 

Invertebrate Zoology: ^™ 

Curator of Insects: A. Neboiss, MSc FRES 
Assistant Curator of Insects: Vacant 
Curator of Invertebrates: B. J. Smith, PhD BSc AMAA 
Assistant Curator of Invertebrates: Suzanne Stevenson BSc 
Assistants: Elizabeth Matheson, Rhyllis J. Plant 

Antk Cuvltov y: oi Anthropology: A. L. West, BA DipSocStud 
Assistant Curator: Alison M. Oates, BA 
Assistants: D. S. Wood, Christine Hogarth, BA 

Survey Officer: 

J. D. Blyth, BSc 

Assistants: R. Houghton, Jane Avery 

Library: . 

Clara Myers, BA, DipLib 

Lorna Crowther, BSc, DipLib 

Assistants: Josie Pearce, Leeanne ODonnell 

Display and Preparation: 

P C. Duce, DipAD, Display Officer 
L. J. Chapman, Senior Preparator 
M. G. Traynor 
B. Hall 
K. Kelly 



S. Huxley, Dip AD 

Diane Stephens, DipArt 

Jan Wheeler ASTC (DipGraphDesign) 

Photographer: 

F. Coffa 

Education Service: 

J. Woollard, TPTC (in charge) 
Brenda Traynor, TITC 
R. L. Miller, DipArt 

G. Wallis, FRMIT (Geol) TSTC 

D. Poad, BA DipEd 
Rosemary Taylor, BA TPTC 
Linda Dunn, TPTC 
Valerie McCormack, (typist) 

Honorary Associates 

with year of appointment 

Geology: 

A. A. Baker, 1951 

A. W. Beasley, MSc PhD DIC, 1973 

A. C. Collins, FRAIA ARIBA AMTPI MACE, 1953 

E. D. Gill, ISM BA BD FGS FRGS, 1973 
Professor J. F. Lovering, MSc PhD, 1974 
J. A. Talent, MSc PhD, 1966 

D. J. Taylor, MSc, 1966 
H. E. Wilkinson, BSc, 1970 

Vertebrate Zoology: 

C. N. Austin, 1955 

C. W. Brazenor, 1962 

A. G. Brown, MRCS (Eng) LRCP (London), 1968 

R. P. Cooper, FNIA, 1952 

N. J. Favaloro, 1945 

A. K. Lee, BSc (WA) MA PhD (Calif), 1972 

M. J. Littlejohn, PhD (WA) MSc, 1972 

P. A. Rawlinson, BSc, 1968 

C. Tanner, 1963 

R. M. Warneke, MSc, BAgrSc, 1966 

H. N. B. Wettenhall, MD BS MRCP FRACP, 1963 

Invertebrate Zoology: 

K. N. Bell, BSc DipEd, 1973 
J. Hope Black, MSc, 1966 
R. F. Burn, 1962 
A. N. Burns, MSc FRES, 1966 

D. F. Crosby, FRES AASA AFAIM, 1968 
R. L. Jensz, BSc DipEd, 1968 

C. McCubbin, 1974 

A. E. Monger, LS MIS (Aust), 1974 

E. T. Smith, 1960 

Jeanette E. Watson, ASMB AMTC, 1970 

L. Winsor, DIP MED TECH FAIMT AAIST, 1974 

Anthropology: 

D. A. Casey, MC FSA, 1933 

J. H. McNamarra, MB BS FRCPA, 1969 
N. M. Wallace, 1970 



CONTENTS 



PAPERS 

PALAEONTOLOGY 



1. A revision of the Gastropod Fauna of the Lily dale Limestone (Early 
Devonian) of Victoria. By C. B. Tassell. (Plates 1-3) 1 

VERTEBRATE ZOOLOGY 

2. A new species of Hemiergis (Scincidae: Lygosominae) from Victoria. By 

A. J. Coventry. 23 

3. The Endemic Australian Lizard Genus Morethia (Scincidae: Lygoso- 
minae) in southern Australia. By P. A. Rawlinson. 27 

INVERTEBRATE ZOOLOGY 

4. A creeping Ctenophoran (Platyctenea: Ctenophora) from Victoria, Aus- 
tralia. By Brian J. Smith and Rhyllis J. Plant. 43 

5. Symbiocladius aurifodinae sp. nov. (Diptera, Chironomidae), A Parasite 
of Nymphs of Australian Leptophlebiidae (Ephemeroptera). By H. B. N. 
Hynes. 47 

6. The Ascidian Fauna of Western Port, Victoria and a comparison with that 

of Port Phillip Bay. By Patricia Kott. 53 

ANTHROPOLOGY 

7. Mortuary Customs of northeast Arnhem Land: an account compiled from 
Donald Thomson's fieldnotes. By Nicolas Peterson. (Plates 4-11) 97 



A REVISION OF THE GASTROPOD FAUNA OF THE LILYDALE 
LIMESTONE (EARLY DEVONIAN) OF VICTORIA 

By C B. Tassell 

* Present Address Albany Residency Museum, Port Road, Albany, W. A. 
Assistant Curator of Fossils, National Museum of Victoria 

Abstract 

Thirteen species of gatsropods are described from the Lower Devonian Lilydale Limestone 
near Melbourne. Two of these are type species of the genera Scafaetrochus (S. lindstrotni 
Etheridge) and Gyrodoma (G. etheridgei (Creswell)). Another two species Michelia brazieri 
(Etheridge) and in part Straparollus {Euomphalus) northi (Etheridge) are the type species 
for two now synonymized genera Vetotuba and Liomphalus. The other species are Tremanotus 
pritchardi Cresswell, Bellerophon (B.) cresswell Etheridge, Phanerotrema australis Etheridge, 
Stenoloron subaequilatera (Chapman), Naticopsis (N.) lilydalensis Cresswell, Murchisonia 
(M.) pritchardi (Etheridge), Siluriphorus antiquus (Cresswell), Loxonema australis (Chap- 
man), and Oriostoma rotundimuratus sp. nov. Also described is Michelia danvhii (de Koninck) 
from the Lower Devonian 'Receptaculites' Limestone, Taemas, New South Wales. The genus 
Boiotremus Horny is considered to be a synonym of Tremanotus Hall. 

The fauna which lacks platyceratids is associated with a diverse coral and stromatoporoid 
assemblage and depleted brachiopod fauna. The gastropod fauna possesses strong affinities 
with the Old World Realm faunas of Europe and North America, and also indicates a 
continuation into the Lower Devonian of certain typically Silurian forms. 



Introduction 

The gastropod fauna of the Lilydale Lime- 
stone was the subject of much attention by 
early palaeontologists in Australia. Robert 
Etheridge, Junr. ? in a series of papers (1890, 
1891, 1894 and 1898) described three new 
genera, Gyrodoma, Scalaetrochus and Veto- 
tuba, nine new species and noted the presence 
of an operculum in place in Straparollus 
{Euomphalus) northi. At about the same time 
the Rev. A. W. Cresswell (1885, 1893 and 
1894), while noting the general elements of 
the fauna described four new species and noted 
the presence of an operculum in place in 5. 
(E.) northi. 

Soon after, Chapman (1916) reviewed the 
entire gastropod fauna. He described one new 
genus Liomphalus, six new species or varieties, 
and noted the presence of Omphalotrochus 
globosum (Schlotheim). Subsequently the fauna 
received very little attention. Knight (1941) 
reviewed the genera established by Etheridge 
and Chapman; Philip and Talent (1959) dis- 
cussed S. (E.) northi and Scalaetrochus lind- 
stromi and more recently Yochelson and Lin- 
sley (1972) discussed the opercula of S. (E.) 
northi and Cyclonema lilydalensis. 

The gastropod fauna of the limestones in the 



Taemas region, N.S.W., has had a similar 
history of early attention and subsequent neg- 
lect. De Koninck (1876) described one new 
genus, Mitchellia, which Knight et al. (p. 1301, 
1960) synonymized with Scoliostoma, seven 
new species and noted the presence of six 
previously described species. 

The basis of this study was the large col- 
lection of the National Museum of Victoria 
which has been collected over a period of 
more than seventy years. The bulk of this has 
come from the limestone where it has been 
the subject of Tertiary weathering (O. P. 
Singleton pers. comm.). Fortunately the wea- 
thering process preferentially destroys the 
matrix of the limestone before destroying the 
fossils. Thus at the right stage in this process 
when the matrix is soft the fossils can be 
obtained free of matrix and moderately well 
preserved. The remainder of the collection is 
preserved in a dense grey limestone. 

Middle Palaeozoic gastropods have been the 
subject of relatively little recent study both in 
Australia and overseas. As a consequence many 
of the genera have been interpreted in a very 
loose sense. So much so, that quite frequently 
authors note that the species they are discussing, 
while being assigned to an established genus 



l 



C. B. TASSELL 



in fact belongs to a new genus. In view of this, 
each of the species from Lilydale has been 
compared with the type species of the appro- 
priate genus. 

However, this has not been done with Cyclo- 
nema australis Etheridge and C. lilydalensis 
Etheridgc, the two species of this genus de- 
scribed from Lilydale. It is generally recognized 
that the Devonian forms assigned to the genus 
Cyclonema in fact constitute at least one new 
genus (Thompson, 1970). To date no one has 
attempted to erect a new genus for these forms 
because a satisfactory basis for distinguishing 
it from Cyclonema sensu stricto is not readily 
apparent. However, the palaeoecological impli- 
cations arising from the assignment of these 
species to the genus Cyclonema are quite 
erroneous. Thus the species from Lilydale are 
not redescribed here, rather it is intended that 
they should be the subject of another study. 
Also not described are members of the classes 
Monoplacophora, Polyplacophora, Pelecypoda, 
and Rostroconchia known to occur at Lilydale. 
These are to be the subject of another paper. 

The generic ranges and distributions given 
by Knight et al. ( 1960) for the genera discussed 
here are accepted in general. Any amendment 
of this is based upon a sensu-sJricto interpreta- 
tion of the genus. 

In this study the following abbreviations 
have been used: P: Palaeontological collection 
of the National Museum of Victoria; F.: 
Australian Museum, Sydney; M.U.G.D.: 
Melbourne University Geology Department; 
A.N.U.: Geology Department, Australian Na- 
tional University. 

All measurements are in millimetres and the 
following symbols relating to the measurements 
have beein used: 
Clu, spiral sculptural elements above the seleni- 

zone. 

Gil, spiral sculptural elements below the selcni- 

zonc. 
Hap, height of aperture. 
Ht, total height of shell. 
L, length measured at the sclenizone in bellero- 

phontids. 
Lap, length of aperture. 
Sw, selenizonc width. 



Wap, width of aperture. 

Wit, width at last trema. 

Wt, total width of shell. 

Wh, total number of whorls in shell. 

*, specimen incomplete. 

Acknowledgements 

I wish to thank Dr A. Ritchie of the Austra- 
lian Museum, Sydney, who made available 
Etheridge's type material; Mr M. Cooper of 
Melbourne University Geology Department, 
for the loan of specimens; Mr T. A. Darragh 
and Dr K. S. W. Campbell for critically reading 
this manuscript and their helpful comments and 
discussion; Dr O. P. Singleton for his discussion 
and comments throughout the course of this 
study; Margaret Tassell for her discussions and 
encouragement and Mr F. Coffa for the photo- 
graphs. 

Age of the Faunas 

In the past, the age of the Lilydale Lime- 
stone has been the subject of some contention 
and frequent revision. Strusz (1972) assessed 
the evidence presently available from a number 
of groups and regarded the Lilydale Limestone 
as being Late Sicgenian in age. 

Strusz (1972) considered the 'Receptaculites' 
Limestone at Taemas, in which Michelia dar- 
wini (de Koninck) occurs, to be Emsian in age. 

However, Philip (1974) summarized recent 
developments in Europe which have placed the 
relationship of the stages in the different facies 
of the Lower Devonian in a state of flux. He 
commented, 'How sterile now seems the de- 
bates as to whether certain limestone horizons 
in eastern Australia are Siegenian or Emsian 
(or even Pragian) in age'. As yet the relation- 
ship of these stages in Europe has not been 
resolved. 

This revision of the gastropod fauna makes 
little contribution to the age determination 
because of the present inadequate knowledge 
of gastropod faunas both in Australia and 
overseas. Most of the genera represented at 
Lilydale are characterized by such long ranges 
that they are of little value in age determina- 
tions. The remaining few genera with relatively 
short ranges such as Scalaetrochus are of limited 
value because of their restricted distribution. 



REVISION OF THE GASTROPOD FAUNA 



Relationships of the Fauna 

Representatives of eight gastropod super- 
families, murchisoniaceans, euomphalaceans, 
bellerophontaceans, pseudophoraceans, pleuro- 
tomariaceans, oriostomataceans, neritaceans 
and loxonemataceans are described from the 
limestone at Lilydale. As well as these there 
are the two species of turbiiniform gastropods 
previously assigned to the genus Cyclonema. 

Comparison with other Lower Devonian 
faunas in Australia is limited by the notable 
lack of recent studies. The closest fauna geo- 
graphically that is described, is that of the 
Marble Creek Limestone (Talent and Philip, 
1956). This fauna is dominated by platycera- 
tids, although species of Tremanotus and 
Michelia are known at Marble Creek as well as 
Lilydale. Chapman (1907, p. 73) also noted 
the presence of Scalaetrochus sp., but this has 
not been subsequently verified. 

The gastropod fauna from the limestones at 
Taemas, N.S.W., as described by de Koninck, 
has only a limited number of genera in com- 
mon with Lilydale. Michelia darwini, described 
here, is very similar in shape to M. brazieri 
from Lilydale. Although de Konimck only de- 
scribed small bellerophontids from Taemas such 
as Bellerophon convolutus de Koninck, large 
forms comparable in size to B. (B.)cresswelli 
Etheridge are known to occur as well. Other- 
wise the gastropod faunas differ considerably, 
with Taemas possessing an abundance of small 
high spired forms, whereas such forms are 
generally absent at Lilydale. 

Boucot (1975) summarized the now con- 
siderable amount of work on Devonian palaeo- 
geography. During t he Lower and Middle 
Devonian a very marked provincialism de- 
veloped when compared with the preceding 
Silurian period and succeeding Late Devonian. 
This provincialism was greatest during the 
Siegenian, Emsian and Eifelian. During the 
Siegenian three realms, Malvinokaffric, Old 
World and Eastern Americas have been recog- 
nized, principally on the basis of brachiopods. 
Other groups including trilobites, corals and 
conodonts also support this biogeographic 
scheme. Within the Old World realm a number 



of regions or sub-provinces are recognized. It 
is within one of these, the Tasman sub-pro- 
vince, that Lilydale is located. 

Boucot has observed that during 'the early 
Devonian the oriostomatids — poleumitids, tre- 
manotids and euomphalids are present only 
in the Old World Realm as "Silurian holdovers" 
(relicts).' They also tend to be particularly 
characteristic of the Bohemian facies of the 
Old World Realm. 

The gastropod fauna at Lilydale strongly 
supports this palaeobiogeographical scheme. 
Fifteen per cent of the fauna consists of relict 
genera Tremanotus, Straparollus (Euomphalus) 
and Oriostoma. These relict genera also tend 
to emphasize the greater similarity between 
Lilydale and the gastropods of the Bohemian 
facies, rather than with the sandier Rhenish 
facies of the Rhenish-Bohemian sub-province. 
The dextrally coiled S. (E.) northi is very like 
the similarly coiled S. (E.) carnicus (Freeh) 
from the Carnac Alps. Also present in the Low- 
er Devonian limestones of the Carnic Alps are 
species of Bellerophon, Phanerotrema and 
Stenoloron which are very similar to the species 
of these genera at Lilydale. Spitz (1907, pi. 13, 
fig. 16 a and b) illustrated a species of Poly- 
tropis from the Carnic Alps which is closely 
similar to the species assigned to 'Cyclonema* 
at Lilydale. Jhaveri (1969) considered that 
the Carnic Alps gastropod fauna shows a close 
relationship with the gastropod faunas from 
the Lower Devonian of Bohemia, Northern 
France and New York as does the fauna 
from Lilydale. 

Palaeoecology 

Two gastropod groups dominate the fauna, 
the murchisoniaceans and the species of 'Cyclo- 
nema'. They constitute about 31% a:nd 25% 
of the fauna respectively. The euomphalaceans, 
bellerophontaceans and pseudophoraceans are 
the next most abundant groups in the fauna 
comprising 13%, 11% and 9% respectively. 
The remaining four superfamilies each com- 
prise about 3% of the fauna. 

In terms of general shape, the turbiniform 
to high spired forms comprise about 70% of 



C. B. TASSELL 



the fauna and the other 30% consists largely of 
discoidal and planispiral forms. About 80% of 
the fauna is cither medium sized or large, 
although this may in part be an artifact of 
selective collecting. However, field comparisons 
of the abundance of smaller forms at Lilydale, 
Buchan and Taemas indicate a very much 
greater abundance at the latter two localities 
than at Lilydale. 

At Lilydale the most distinctive feature of 
the fauna is the total absence of members of 
the Family Platyccratidae which is presently 
characterized by a cophrophagous mode of 
life. The two species of 'Cyclonema* described 
from Lilydale possess an operculum (Yochel- 
son and Linsley, 1972), lack any apertural or 
coiling irregularity and possess no other feature 
that could be suggestive of a cophrophagous 
mode of life. Thus their association with the 
platyccratidae would greatly distort any inter- 
pretation of the environment of the fauna. 

Not only is there a general absence of platy- 
ceratids at Lilydale, there is also a general lack 
of crinoidal remains in the limestone. A signifi- 
cant percentage of the crinoidol remains present 
arc composed of fragments of Pcmerocrinus 
(Bates, 1972). This is in marked contrast to 
the gastropod fauna of the Marble Creek lime- 
stone. This latter fauna, dominated by platy- 
ccratids, occurs in a limestone composed in 
large part of crinoidal fragments. The fauna is 
also notable for its low taxomomic diversity in 
comparison with the diversity at Lilydale. 

A similar situation is seen in the diversity of 
the gastropod fauna in the Silurian recfal com- 
plexes of Gotland, Sweden. The reefal lime- 
stones in which Manten (1971) observed 
gastropod faunas had a considerably more 
diverse fauna than that of the associated cri- 
noidal limestones. 

Linsley (1968) in his description of the 
Middle Devonian gastropods of the Anderdon 
Limestone recognized two principal habitats. 
One of these is a 'biostromal' environment 
where small snails lived on the carbonate mud 
flats between the corals and stromatoporoids in 
relatively shallow water. The gastropods of the 
other habitat, that of the 'inter-reef lived on 
the carbonate muds developed between sparse, 



localized tetracoral assemblages. This fauna 
was distinguished by its considerably larger 
size, l"-5" being the maximum dimension. Also 
present were a few nautiloids, articulate 
brachiopods and some ostracodes. Linsley 
considered that the fauna and the sedimeait 
suggested a fairly quiet environment. 

The composition of the fauna of Lilydale 
resembles reasonably closely the fauna of Les- 
ley's inter-reef habitat. Both are dominated by 
large forms although more of these are high- 
spired types at Lilydale. However, the first spec- 
ies of Scalaeirochus to be recorded outside 
south-eastern Australia was described by Lins- 
ley from the Anderdon Limestone. Thus the 
similarity of sediment and fauna suggest that 
the gastropods of the Lilydale Limestone oc- 
cupied an environment somewhat similar to 
the inter-reef environment of the Anderdon 
Limestone. 

Yochelson and Dutro (1960) in their de- 
scription of a Mississippian and Permian 
gastropod fauna from limestones in northern 
Alaska also made some comments on the 
palaeoecology of the assemblages. They ob- 
served that 'Platyceras commonly occurs here 
in crinoidal limestones'. This is in accord- 
ance with its distribution in the limestones at 
Gotland, Marble Creek and Lilydale. Where 
Platyceras is common m the Lower Mississip- 
pian sediments, 'the associated gastropods show 
less variety than in the Upper Mississippian' 
(where Platyceras is less common). Again this 
is comparable with the situation at Lilydale 
and Gotland. 

However, they noted that 'corals and gastro- 
pods also appear to be nearly mutually exclu- 
sive' and that 'gastropods are commonly as- 
sociated with numerous taxonomically diver- 
sified brachiopods'. At Lilydale a situation 
quite the reverse exists. Gastropods and corals 
are very closely associated while only a de- 
pleted and restricted brachiopod fauna is pre- 
sent. 

That such diverse gastropod associations 
exist only serves to emphasize the need for 
further work on this group before more mean- 
ingful generalizations on gastropod palaeoe- 
cology can be made. 



REVISION OF THE GASTROPOD FAUNA 



Systematic Descriptions 

Superfamily BELLEROPHONTACEA 

McCoy, 1851 
Family Sinuitidae Dall in Zittel-Eastman, 

1913 
Subfamily Tremanotinae Peel, 1972 
Genus Tremanotus Hall, 1865 

( = Boiotremus Horny, 1962). 

Type Species; Tremanotus alpheus Hall, 1865; 
Middle Silurian; Bridgeport, Illinois, U.S.A. 
Range: Middle Ordovician to Lower Devonian. 
The presence of 7\ pritchardi at Lilydale and 
T. fortis Freeh and T. insectus Freeh in the 
Upper Koneprusy Limestone, Koneprusy ex- 
tends the upper range of the genus from Middle 
Silurian to Lower Devonian. 
Discussion: Knight et al. (1960) provided a 
diagnosis for this genus in which the slit is 
'represented by a row of tremata, all but the 
last few closed, not extending on to expanded 
lip'. Subsequently Horny (1963) amended this 
diagnosis to read, 'slit represented by a row of 
tremata in body whorls, not extending on the 
expanded lip; no tremata but shallow sinus in 
outer lip in young stages'. Horny (1962) also 
erected a new genus, Boiotremus, characterized 
by 'tremata present along the whole length of 
the whorls, periodical widened apertures after 
distances of 1-3 tremata'. It is into this latter 
genus that most of the species previously as- 
signed to Tremanotus would be placed. 

In his discussion of the genus Tremanotus 
he noted that the 'main and characteristic sign 
is the existence of the five opened tremata in 
the body whorl region. In the ontogenetically 
younger stage there are no tremata developed 
, . .' This interpretation is inconsistent with T. 
alpheus as described by Hall and Knight's 
(1941) redescription which Horny (1963, p. 
97) suggested implicitly supported his case. 
Knight (noted '6-8 tremata remaining open' and 
'the earlier ones (were) filled'. That there are 
more than 5 tremata can be clearly seen in 
Knight's figures of the type species. The figures 
of Clarke and Ruedemann (1903) also clearly 
indicate that the presence of tremata is not 
confined to the body whorl. Unfortunately the 
preservation and orientation of the specimens 



figured by Horny does not show the dorsal 
surfaces of the earlier whorls. 

Thus Horny's amendment of the diagnosis 
of Tremanotus is based upon a misconception 
and is without justification. The new genus 
Boiotremus to which he attributed 'all tre- 
manotids' which have developed tremata during 
the whole life of the specimen, i.e. in all onto- 
genie stages' is thus a synonym of Tremanotus. 

Horny (1963, p. 97) in his discussion of 
Tremanotus as redefined by himself, considered 
that both the tremata and the flared aperture 
were features which developed only at maturity. 
However, the development of the tremata was 
dependent upon the development of the flared 
aperture. This concept of the growth sequence 
with its inherent reorganization of the exhalent 
system is without justification because of the 
presence of tremata throughout the develop- 
ment of the entire shell. 

Tremanotus pritchardi Cresswell, 1893 
(PL 1, fig. 17) 

1893 Tremanotus pritchardi Cresswell, p. 42, pi. 8, 

fig. 1. 
1913 Tremanotus pritchardi Cresswell; Chapman, p. 

227. 
1916 Tremanotus pritchardi Cresswell; Chapman, p. 

79 in part. 

Diagnosis: Large form of genus in which the 
relationship between major growth rugae and 
tremata is variable; numerous fine growth lines 
a«nd open tremata are present. 
Description: Large planispiral gastropod with 
a widely expanded aperture in the final growth 
stage; wide umbilici; whorl profile gently arched 
dorsally, more strongly curved on the sides 
turning sharply into the wide and deep umbilici, 
flattened on the inner surface; aperture in final 
growth stage sub-oval; neither a sinus nor 
tremata are developed on the dorsal surface 
of the expanded region of the aperture; pos- 
terior to the expanded apertural region, the 
existence of a small sinus situated medially on 
the dorsal crest of the whorl is indicated by a 
slight posterior flexure of the growth lines; along 
the resultant selenizone numerous ovoid tre- 
mata are developed, the most anterior trema 
is represented by a solid protrusion over which 
growth lines pass; then follows a number of 
open tremata, up to 11, the tremata preceding 



C. B. TASSELL 



the open trcmata are sealed and flush with the 
whorl surface; beween tremata the growth lines 
are directed posteriorly towards the earlier 
tremata; as well as the closely spaced growth 
lines, prominent growth rugae are developed; 
the relationship of the growth rugae to the 
tremata is variable; in some cases the rugae 
intersect the selenizone between trcmata, in 
others at the trema; sculpture is composed of 
numerous spiral costae which arise in the 
umbilici; sculptural elements vary from one to 
three orders, sculpture and growth lines form 
a reticulate pattern over the entire whorl sur- 
face. 



Dimensions: 


L 


Wt 


Lap 


Wap 


Wit 


Wh 


M.U.G.D. 1666 
P914 


92 


56 

47 


46 
43 


43 
40 


35 
33 


5+ 



Location of Types: Melbourne University Geo- 
logy Department. Holotype, M.U.G.D. 1666 
and counterpart M.U.G.D. 1667, G. B. Prit- 
chard Coll. 

Material: Holotype, counterpart, and 12 other 
specimens. 

Discussion: This species is distinguished from 
the type species, T. alpheus, by having more 
open tremata, up to 11 as compared to the 
latter's 6 to 8. Transverse growth lines, princi- 
pally fine ones of a type lacking in the American 
form, are more abundant. The relationship of 
the growth rugae and the tremata is quite 
variable in the Lilydale form. However, rugae 
only occur between tremata in the type species. 
The last potential trema is closed, whereas no 
mention of this was made by Knight (1941, 
p. 354) in his redescription of the type species. 
Knight postulated that the tremata were 
formed during periods of growth when the 
flared aperture was not being deposited. At 
such times a shallow sinus in the outer dorsal 
lip formed a short slit, subsequently closed 
anteriorly by growth of the flared aperture. 
The flared aperture was then resorbed, the pro- 
minent growth rugae marking the point to 
which resorption occurred. This was followed 
by the initial growth type in which another 
trema was formed. This cycle was repeated 
numerous times as is indicated by the number 
of tremata. 



Material from Lilydale suggests a mode of 
growth in T. pritchardi different from that 
postulated by Knight. 

Initial growth, as in Haliotis, saw the de- 
velopment of a short sinus, which with subse- 
quent growth of the apertural lips was closed, 
so forming a trema. Then followed a period in 
which only an exceedingly narrow and shallow 
sinus was developed. Subsequently this sinus 
deepened and another trema formed. With 
growth, earlier tremata were no longer required 
and sealed as in Haliotis. This process of growth 
continued until the animal reached its penulti- 
mate growth stage. At this point the mode of 
growth changed. No longer was a sinus formed. 
Rather the prominent flared aperture was de- 
veloped. Frequently the sculpture present 
on the flared aperture is quite different to the 
rest of the shell, e.g. all but the first order spiral 
sculpture may be absent. 

This growth cycle differs from that suggested 
by Knight, in that it does not require the flared 

PLATE 1 

Fig. 1 — Michelia danvini (de Koninck), ANU 

36852, hypotvpe, Bloomfield Property, Yass, 
N.SAV., XI. 

Fig. 2 —Michelia darwini (de Koninck), ANU 

36853, hypotvpe, Bloomfield Property, Yass, 
N.S.W., XL 

Michelia brazieri (Etheridge), P1058, hypo- 
type, XI. 

— Loxonema australis (Chapman), P38508, 
hypotype, XI. 

Michelia brazieri (Etheridge), P1059, hypo- 
type, XI. 

— Belterophon (Bellerophon) cresswelli Eth- 
eridge, P12838, hypotype, X 2/3. 
•Straparollus {Euomphalus) northi (Eth- 
eridge). P28716. hypotype, X 21. 

— Straparollus {Euomphalus) northi (Eth- 
eridge), P28714, hypotype, X 2/3. 

10 — Naticopsis (Naticopsis) lihdalensis Cress- 
well, P37740, hypotype, XT. 

1-12 — Phanerotrema australis Etheridge, F. 
1332, syntype, X H (approx.). 

— Scalaetrochus lindstromi Etheridge, F.1137, 
holotype, X 2/3. 

— Stcnoloron subaequ'tlatera (Chapman), 
P37643, hypotype, X 2/3. 

: — Scalaetrochus lindstromi Etheridge, F.1137, 
holotype X 2/3. 

Scalaetrochus lindstromi Etheridge, P38505, 
hypotype X 2/3. Basal view showing open 
umbilicus. 
Fig. 17 — Tremanotus pritchardi Cresswell, M.U.G.D. 

1666, holotype, X 2/3. 
Fig. 18 — Scalaetrochus lindstromi Etheridge, F.1137, 
holotype, X 2/3. Basal view. 



Fig. 


3 • 


Fig. 


4 • 


Fig. 


5 - 


Fig. 


6 ■ 


Fig. 


7 - 


Fig. 


8 ■ 


Figs 


. 9- 


Figs 


. 1 


Fig. 


13- 


Fig. 


14- 


Fig. 


15- 


Fig. 


16- 



MEM. NAT. MUS. VICT. 37 PLATE 1 





;,' 






^ 




5 


















REVISION OF THE GASTROPOD FAUNA 



aperture to be deposited and resorbed numerous 
times. Here the flared aperture is considered 
to be a gerontic feature. It is also considered 
that the type species mode of growth was the 
same as that of T. pritchardL 

In his discussion of Boiotremus, Horny 
(1963, p. 101) noted the presence of tremata 
throughout the entire life of the specimen and 
the development of periodically widened aper- 
tures. He did not describe the extent and 
nature of this 'periodical widened aperture'. 
That Tremanotus did not develop a large 
flared aperture periodically has been discussed 
earlier. Horny did not make mention of resorp- 
tion of this widened aperture. The only other 
method of removal would be purely mechanical 
by abrasion. Whatever the method, the removal 
of this region results in the formation of a 
feature he termed a 'scar 1 . This is in fact the 
same as the major growth rugae observed in 
the type species and the species from Lilydale. 
As previously noted the distribution and de- 
gree of development of these growth rugae 
is quite variable amongst members of the same 
species and between different species. This is 
also true of the species figured by Horny. 

Although Horny (1963) figured two speci- 
mens NM-L5727 and NM-L5729 (PI. 26, 
fig. 2 and PI. 27, figs. 2, 3, 4 and 5) which he 
considered show the widened aperture, these 
specimens could equally as well be mature 
specimens with badly damaged expanded aper- 
tural regions. Because of the great variability 
in the development of the growth rugae or 
'scars' it is considered highly unlikely that the 
widened apertures as described by Horny were 
developed. To demonstrate their existence un- 
equivocally would require a mature specimen 
with both the final expanded apcrtural region 
and the older widened apertures preserved. 
Their appearance would be much the same as 
varices of certain gastropods. If, in fact they do 
exist, their great variability as reflected by the 
growth rugae or 'scars' would make them un- 
satisfactory as a generic characteristic. 

Chapman (1916, p. 79) mentioned a speci- 
men from Marble Creek, Thomson River, 
Victoria. Subsequently Talent and Philip (1956) 
described a new species T. cyclocostatus from 



this locality. This is distinguished from T. 
pritchardi by its considerably smaller size and 
much finer growth lines and sculpture. It also 
has fewer foliaceous growth rugae and those 
that are present are irregularly developed. 
Subfamily Bellerophontinae McCoy, 1851. 

Genus Bellerophon Montford, 1808 

Subgenus Bellerophon (Bellerophon) Mont- 
fort, 1808 

Type Species: Bellerophom vasulites Montfort, 
1808; Middle Devonian; The Eifel, Germany. 

Bellerophon (Bellerophon) cresswelli Etheridge, 
1891 

(PI. 1, fig. 6; PI. 3, figs. 3, 4, 5, 6, 9) 

1891 Bellerophon cresswelli Etheridgc, p. 130, pi. 19, 

figs. 6-8. 
1913 Bellerophon cresswelli Etheridge; Chapman, p. 

227. 
1916 Bellerophon cresswelli Etheridge; Chapman, p. 

80, pi. 2, fig. 12, pi. 4, fig. 53. 
1916 Bellerophon pisutn Chapman, p. 80 pi. 2, figs. 

9-11. 

Diagnosis; Typical form of genus with thick- 
ened outer lip which extends posteriorly beyond 
the umbilical region; broadly crescentic aper- 
ture; selenizone only slightly elevated and bor- 
dered by two fine threads; sculpture composed 
of very line transverse elements. 
Description: Medium, subglobular, narrowly 
umbilicate planispiral gastropod with broad 
involute whorls; whorl profile gently arched 
dorsally, more strongly curved on the sides, 
turning sharply into the narrow umbilici; aper- 
ture very broadly crescentic; margin of outer 
lip not flared anteriorly but flared outward in 
the lateral and umbilical regions, margin con- 
tinues across the parietal wall as a moderately 
thick inductura which thickens considerably 
towards the lateral margins, greatest thickness 
at the junction of the flared outer lip and 
parietal wall; narrow, moderately deep slit on 
outer lip generating a dorsal selenizone which 
is very slightly raised above the whorl surface 
and bordered by two very fine threads; sculpture 
consists of fine transvere growth lines with 
occasional growth rugae, holotype has about 
seven whorls. 



C. B. TASSELL 



Dimensions: 



Wt 



Lap Wap Wh Sw 



F.1327 
P293 

P1087 
P12837 
P12838 
P34938 



48 
11 
41 
44 
12 



37 
47 
10 
44 
43 
12 



29 

23 

3* 
37 
37 

3 



25 
41 

7-3 
40 
25* 
9# 



7 1 
6 

— 4 

— 1 

— 4 



Location of Types; 1. B. cresswelli, Australian 
Museum, Holotype, F.1327. National Museum 
of Victoria, Hypotypes, PI 2838, P34938. G. 
Sweet Coll. 

2. B. pisum, National Museum of Victoria, 
Holotype, P1087, A. W. Cresswell Coll. 
Material: Holotype, 3 hypotypes and 37 other 
specimens. 

Discussion: The holotype is partially broken, 
revealing the inner whorls which number about 
seven. This species is distinguishable from the 
similar sized type species B. (B.) vasulites by a 
number of features. The aperture of B. (B.) 
cresswelli is more broadly crescentic in shape. 
The outer lip is considerably thicker in the 
Lilydale form and extends further posteriorly 
beyond the umbilici. The selenizonc of the 
type species is raised further above the whorl 
surface and bordered by shallow depressions 
rather than the fine threads of the Lilydale 
form. Transverse sculpture in B. (B.) cress- 
welli is considerably finer, particularly in the 
regions near the selenizone. 

Chapman (1916, p. 80) referred to a well 
preserved specimen in the National Museum 
of Victoria collection which exhibits 'a faint 
but definite lattice structure of wavy striae 
across the growth lines'. However, none of the 
specimens examined in this study possess such 
ornamentation. 

During growth B. (B.) cresswelli changed 
slightly in form. The aperture became more 
flared. In the juvenile form, flaring was con- 
fined to the umbilical region, but with growth 
the lateral areas of the outer lip also became 
flared. Initially quite thin, the inductura also 
thickened with growth, as did the outer lip. 
The influence of thickening is quite marked 
in the change of shape of the junctions of the 
parietal inductura and outer lip. In the juvenile 
forms, a marked, moderately deep channel is 
present, whereas in the older forms the channel 



is considerably shallower. The sculpture also 
changes from being quite wide, regular and 
nearly foliaceous in smaller forms to generally 
closer, finer but more irregular and variable in 
the larger forms. 

Chapman (1916, p. 80) described B. (B.) 
pisum from Lilydale. He did not indicate spe- 
cifically how it is distinguishable from this 
species. Presumably it is because of its smaller 
size, distinct sinus in the outer lip and sculpture 
of 'interrupted radial striae . . . between the 
lines of growth'. However, none of the features 
that Chapman mentioned in his description of 
B. pisum are unique. All these features are 
found in specimens of B, (B.) cresswelli, par- 
ticularly the smaller forms. It is considered that 
B. pisum is a juvenile of B. (B.) cresswelli. 

Superfamily EUOMPHALACEA de Koninck, 
1881 

Family Euomphalidae de Koninck, 1881 

Genus Straparollus Montfort, 1810 

Subgenus Straparollus (Euomphalus) J. 

Sowerby, 1814 

(= Liomphalus Chapman, 1916) 
Type Species: Euomphalus pentangulatus J. 
Sowerby, 1814; Lower Carboniferous; near 
Dublin, Ireland. 

Discussion; Chapman (1916, p. 90) erected the 
genus Liomphalus which he distinguished from 
Euomphalus in having 'smooth rounded unian- 
gulated whorls', and from Straparollus in 
possessing a concave spire. The genus is 
characterized by the following: discoidal; base 
concave; wide umbilicus; spire depressed; 
whorls smooth, sometimes with keel; whorls 
thicken progressively, in the late stages free 
or adpressed. 

Knight (1941, p. 174) considered it 'utterly 
impossible to arrive at any comprehension of 
the genotype species except by comparison 
with the adequately described species of other 
authors referred by Chapman to his genus'. 
Later, Knight (1944, p. 465) placed it in 
synonomy with Lytospira Koken. 

Philip and Talent (1959, p. 50) demon- 
strated that the genus Liomphalus is based 
on the internal moulds of Straparollus {Euom- 
phalus) northi (Etheridge). Their conclusions 



REVISION OF THE GASTROPOD FAUNA 



are amply supported by many specimens in the 
collections studied. 

Straparollus (Euomphalus) northi (Etheridge), 

1890 

(PI. 1, figs. 7, 8. PL 2, fig. 11. PL 3, figs. 

1, 2, 7, 8) 

1890 Oriostoma northi Etheridge, p. 64, pi. 9, figs. 

6-7. 
1894 Oriostoma northi Etheridge, p. 151, pi. 9, 

figs. 1-4. 
1894 Euomphalus (Oriostoma) northi Etheridge; 

Cresswell, p. 157. 
1913 Euomphalus northi (Etheridge); Chapman, p. 

227. 
1916 Euomphalus northi (Etheridge); Chapman, 

p. 90. 
1916 Liomphalus australis Chapman, p. 90, pi. 4, 

figs. 32-33. 
1959 Straparolus (Euomphalus) northi (Etheridge); 

Philip and Talent, p. 50, pi. 7, figs. 1-12, 

pi. 8, figs. 1-2. 
1972 Oriostoma northi Etheridge; Yochelson and 

Linsley, p. 8, pi. 1, fig. 6, pi. 2, figs. 1-5. 

Diagnosis: Dextrally coiled discoidal gastro- 
pod; variably developed angulations on upper 
and lower whorl surfaces; sculpture initially 
consists of strong transverse costae becoming 
less developed with growth. 
Description: Medium to large discoidal dextral 
gastropod; numerous whorls with profile flat to 
gently convex and inward sloping between the 
upper suture and the variably developed upper 
keel at the junction of the upper and outer 
whorl surfaces; more than one keel may be 
developed on the upper surface; the prominence 
of the keel or keels tends to decrease with 
increased size; the junction of the outer and 
basal whorl surfaces is generally less pro- 
nounced than that for the upper surfaces; 
sutures impressed; base strongly arched; very 
wide umbilicus; spire depressed; aperture 
circular; columellar lip thin, concave parietal 
lip thin; outer lip slightly thicker and it extends 
outwards radially from the upper suture to the 
outer whorl surface where a mild concave 
flexure is sometimes developed, and then con- 
tinues inwards across the base radially to the 
suture; no sinus or flexure of growth 
lines is developed at the keel, if pre- 
sent; sculpture is variably developed on 
specimens of different size and on the one 
specimen; the smaller specimens and the inner 
whorls of the larger specimens possess promi- 



nent transverse ridges; on the larger specimens, 
fine growth lines gradually succeed the promi- 
nent juvenile sculpture; sculpture on the base 
while similar to that on the upper and outer 
surfaces is less strongly developed; transverse 
partitions occur in early whorls; multispiral 
operculum of numerous fine whorls of variable 
thickness, circular in shape; whorls are normally 
visible on the plano-concave exterior surface; 
the concave internal surface is smooth being 
composed of laminae deposited nearly at right 
angles to the opercula rim; the degree to which 
the internal surface is concave is quite variable; 
a shallow central depression occupies about 
one-third of the inner surface; thickness is 
variable, even for opercula of the same dia- 
meter; operculum fits tightly in the aperture, 
being retracted into the shell for about 2 mm. 



Dimensions: 














Ht 


Wt 


Hap 


Wap 


Wh 


F.1321a 


135 


74 


— 


— 


4 


F.1139e 


20-2 


51 


— 


— 


— 


P1107 


25* 


71 


23 


23 


4 


P2503 


— 


54 


— 


— 


3 


P7609 


— 


60 


— 


— 


— 


P28373 


32 


97 


— 


— 


— 


P28498 


19 


63 


— 


— 


4+ 


P28499 


— 


59 


— 


— 




P28707 


14 


49 


— 


— 


3 + 


P28711 


25 


77 


— 


— 


4+ 


P28712 


27 


77 


— 


— 


3 + 


P28714 


16 


44 


— 


— 


4 


P28716 


10 


19 


— 


— 


4 


P28719 


7 


14 


— 


— 


— 


P343O0 


27 


54 


— 


— 


4 



Location of Types: 1. Oriostoma northi, Aus- 
tralian Museum. Holotype, F. 1321a, Paratype, 
F. 1 139e. National Museum of Victoria. Hypo- 
types, P1107, P1115. A. W. Cresswell Coll. 
P26890, P28499, P28714, P28716, P28718, 
P28719. E. D. Gill Coll. and P34300-34302 
which were formerly GSV 55329, 55330 and 
55332. 

2. Liomphalus australis, National Museum 
of Victoria. Holotype, P7609, Paratype, P2503. 
A. W. Cresswell Coll. 

Material: Holotype, paratype, 13 hypotypes and 
49 other specimens. 

Discussion: Comparison of S. (£.) northi with 
the type species S. (£.) pentangulatus reveals 
a number of differences. The former has a 
shallower, wider umbilicus and is coiled dex- 



10 



C. B. TASSELL 



trally. Its upper keel is less strongly developed 
and there is an absence of spiral sculptural 
elements. The sculpture of the Lilydale form 
is more variable; initially the transverse ele- 
ments are much stronger than those of the type 
species but finally they are considerably weaker. 
The sculpture is also less evenly developed on 
the upper and basal whorl surfaces. 

The dextral coiling of S. (E.) northi is also 
a feature of S. (E.) carnicus (Freeh). As 
figured by Jhaveri (1969, pi. 21, fig. 8) this 
species also possesses strong transverse sculp- 
tural elements as does S. (E.) northi. It also 
probably possesses an operculum which is very 
similar to that of S. (E.) northi, (Yochelson 
and Linsley 1972, p. 9). 

Yochelson and Linsley (1972, p. 8) de- 
scribed the operculum from S. (E.) northi and 
compared it with very similar types found in 
a number of other genera which belong to more 
than one family. Because of this they suggested 
implicitly that a revision of these families was 
needed. Thus they preferred to leave the species 
from Lilydale in the genus Oriostoma as 
originally determined by Etheridge. However, 
comparison of S. (E.) northi with Oriostoma 
barrandei Munier-Chalmas, the type species 
reveals sufficient differences to preclude it from 
belonging to the latter genus. S. (E.) northi 
is discoidal rather than turbiniform and has a 
much wider umbilicus. There is an absence of 
the spiral sculpture characteristic of the type 
species. The form from Lilydale has a more 
angular whorl profile and its aperture is more 
circular in shape. 

Superfamily PLEUROTOMARIACEA Swain- 
son, 1840 

Family Phanerotrematidae Knight, 

1956 

Genus Phanerotrema Fischer, 1885 

Type Species: Pleurotomaria labrosa Hall, 
1860; Lower Devonian; Carlisle, New York, 
United States of America. 

Phanerotrema australis Etheridge, 1891 

(PL l,figs. 11, 12. PI. 2, figs. 3, 10,12) 

1891 Phanerotrema australis Etheridge, p. 128, pi. 
19, figs. 4-5. 



1913 Phanerotrema australis Etheridge; Chapman, p. 

227. 
1916 Phanerotrema australis Etheridge; Chapman, p. 

83, pi. 3, fig. 25. 
Diagnosis: Typical form of genus with thick, 
short, straight columellar lip, well-developed 
parietal inductura and simple rectangular sculp- 
ture pattern arising from the intersection of 
the collabral growth lines and spiral cords. 
Description: Large, turbiniform gastropod with 
few whorls; whorl profile sub-angular, gently 
arched above and below the selenizone at the 
angular periphery; periphery high above mid- 
whorl; sutures deeply impressed to sub-canali- 
culate; body whorl greatly expanded; umbilicus 
absent; columellar lip thickened, continuous 
with the thick extensive parietal inductura; 
outer lip thin with a broad sinus that forms a 
shallow slit at the periphery which gives rise 
to the selenizone; from the upper suture to the 
selenizone outer lip very gently prosocline; 
below the selenizone the outer lip is gently 
prosocline; gently concave selenizone mod- 
erately wide, and bordered by two threads; 
fine collabral growth lines and infrequent 
growth rugae, cancellated by two orders of 
fine spiral cords to form a rectangular pattern 
over the entire whorl surface, occasional speci- 
mens have a retrousse intersection. 

PLATE 2 

-Michelia brazieri (Etheridge), F.1145, 

holotype, XI (approx.). 
-Gyrodoma etheridgei (Cresswell), F.2542, 

hypotype, XI (approx.). 
-Phanerotrema australis Etheridge, P.41706, 

hypotype, X 2/3. 
-Siluriphorus antiquus (Cresswell) , P918, 

hypotype, XI. Oblique basal view. 
-Scalaetrochus lindstromi Etheridge, P39279, 

hypotype, X 11 (approx.). Basal view show- 
ing prominent peripheral frill. 
-Siluriphorus antiquus (Cresswell), P917, 

holotype, XI. Apical view. 
-Oriostoma rotundimuratus sp. nov., P1089, 

holotype, Apical view. 
-Loxonema australis (Chapman), P12851, 

holotype, X li. 
-Gyrodoma etheridgei (Cresswell), P10187, 

holotype, X 1. 
-Phanerotrema australis Etheridge, F.39308, 

syntype, X 2/3. 
-Straparollus (Euomphalus) northi (Eth- 
eridge), F.1321a, holotype, X 2/3. Apical 

view. 
-Phanerotrema australis Etheridge, F.39308, 

syntype, X 2/3. 



Fig. 


1 


Fig. 


2 . 


Fig. 


3 


Fig. 


4 


Fig. 


5 • 


Fig. 


6 


Fig. 


7 • 


Fig. 


8 ■ 


Fig. 


9 


Fig. 


10 


Fig. 


11 



Fig. 12- 



MEM. NAT. MUS. VICT. 37 PLATE 2 



v.'/"":"- 



W 1 







'" ' *\ 



) 



'■ 



:-::,.?:,■:■> 



&%§&n > 



>, 












i» 



Sk 1 : 



to 






8 





9 







v 



; . 



12 



REVISION OF THE GASTROPOD FAUNA 



11 



Dimensions: 

Ht Wt Hap Wap Wh Clu Cll 

F.1332 24 21 — — 4 14 16+ 

F.39308 82 73 — — 24- — — 

P384 63* 47 — — 3+ 29 31 

P12841 93 78 — — 4 28* 28* 

Location of Types: Australian Museum. Syn- 
types, R1332 and F.39308. National Museum 
of Victoria. Hypotypes, P 12841, presented by 
Dr E. Brooke Nicholls and P41706, A. W. 
Cresswell Coll. 

Material: Two syntypes, 2 hypotypes and 12 
other specimens. 

Discussion: P. Australis differs from the type 
species in having a straighter, more thickened 
and longer columellar lip. The type species' 
columellar lip is markedly curved. The parietal 
inductura on the form from Lilydale is also 
thicker than that of the type species. The 
intersection of the collabral growth lines and 
spiral sculptural elements in the type species 
is retrousse, whereas that of the Lilydale form 
is generally considerably simpler. 

Family Gosseletinidae Wenz, 1938 

Genus Stenoloron Oehlert, 1888 
Type Species: Pleurotomaria viennayi Oehlert, 
1888; Lower Devonian; Saint-Roch (La Bacon- 
nier), department de la Mayenne, France. 
Discussion: The presence at Lilydale of a 
member of this genus extends the known dis- 
tribution as it was previously confined to 
Europe and North America. 

Stenoloron subaequilatera (Chapman), 1916 

(PL 1, fig. 14) 

1916 Mourlonia subaequilatera Chapman, p. 83, pi, 
3, figs. 18-19. 

Diagnosis: Typical form of genus with a seleni- 

zone bordered by two cords close to mid-whorl 

periphery and finely developed spiral elements 

of sculpture. 

Description: Medium rotelliform, umbilicate 

gastropod; whorl profile well rounded, convex; 

moderately impressed sutures; base rounded; 

aperture known only in part; outer lip with a 

moderately deep and angular sinus that forms 

a slit which gives rise to a narrow selenizone; 

selenizonc located about one-third of the way 

between the mid-whorl periphery and upper 

suture; between the upper suture and the seleni- 



zone the outer lip is prosocline with a moderate 
obliquity; below the selenizone it is prosocyrt, 
passing forwards for a short distance before 
rounding gently and passing nearly radially 
across the base; inner lip not known; seleni- 
zone depressed and bordered by two moderately 
developed cords; collabral lines strongly de- 
veloped; very subdued elements of spiral sculp- 
ture. 



Dimensions: 



P925 
P37643 



H t Wt Ha p Wap Wh S w 

— 5 0-4 

— 6 8 



16 

56 



22 5 

61 — — 6 

Location of Types: National Museum of Vic- 
toria. Holotype, P925, A. W. Cresswell Coll. 
Hypotypc, P37643. 

Material: Holotype, hypotype and 2 other 
specimens. 

Discussion: S. subaequilatera is represented by 
only a few specimens, none of which is com- 
plete. However, it is possible to distinguish it 
from the type species as known on the basis 
of Oehlert's original figures and description. 
The type species' selenizone is located about 
mid-way between the periphery and upper 
suture whereas that of the Lilydale form is 
considerably closer to the periphery. The form 
from Lilydale also possesses two cords border- 
ing the selenizone and fine spiral elements of 
sculpture. As figured by Oehlert (1888, pi. 9, 
figs. 2 and 2a) it is quite possible that the cords 
are present although not mentioned in 
the description. However, there is no sug- 
gestion in Oehlert's figures of the presence of 
fine spiral elements of sculpture. 

Superfamily ORIOSTOMATACEA Wenz, 

1938 
Family Ortostomatidae Wenz, 1938 

Genus Oriostoma Munier-Chalmas, 1876 

Type Species: Oriostoma barrandei Munier- 
Chalmas, 1876; Lower Devonian; Bois Roux 
quarry at Gahard, near Rennes, France. 

Oriostoma rotundimuratus sp. nov. 

(PI. 2, fig. 7) 
1916 Omphalotrochus zlobosum (Schlotheim); Chap- 
man, p. 92, pi. 4, figs. 35-36. 

Diagnosis: Small form of genus very similar 



12 



C. B. TASSELL 



to the type species but with stronger sculpture 
and more subdued collabral lines. 
Description: Small, low spired, turbiniform gas- 
tropod with a few whorls in slight contact; 
outer whorl frequently disjunct; whorls increase 
rapidly in size, body whorl large; whorl 
profile rounded, arching upwards from 
the upper suture to the sub-rounded 
shoulder, then arching more gently to the 
lower slightly less rounded basal angulation 
surrounding the umbilicus, finally passing in 
a gentle convex arch to the umbilicus; sutures 
deep; umbilicate; columellar lip thin; parietal 
lip thin; outer lip thick, weakly prosocline; 
without sinus or slit; retrousse at the shoulder, 
and at each of the elements of spiral sculpture; 
sculpture composed of a number of strong 
spiral elements of at least two orders; collabral 
growth lines fine, slightly foliaceous and 
retrousse over each of the spiral sculptural 
elements. 



Dimensions: 



Ht 



Wt Hap Wap Wh 



12 3 

7 — — 3 



P1089 7 

P12850 7 

P37644 5 

Location of Types: National Museum of Vic- 
toria. Holotype, P1089; hypotype, P1088. 
A. W. Cresswell Coll. 

Material: Holotype, hypotype and 7 other 
specimens. 

Discussion: Comparison of O. rotundimuratus 
with the type species as redescribed by Knight 
(1941, p. 219) reveals few differences. The 
form from Lilydale has a more rounded whorl 
profile particularly in the region of the shoulder. 
While the collabral lines of the type species are 
more prominent, the spiral elements are 
weaker. The type species also possesses a more 
arcuate columellar lip and is slightly larger. 

Chapman (1916, p. 93) considered this 
species to be Trochilites globosus Schlotheim 
(1820, p. 162). Comparison of the Lilydale 
form with this species is rather difficult, for 
Schlotheim's description is brief and he pro- 
vided no figures. However, Lindstrom (1884, 
p. 162) studied the original specimen from 
Gotland upon which Schlotheim based his des- 
cription. He synonymized Euomphalus funatus 



Sowerby (1823, p. 71) with it. 

Comparison of the Lilydale form with illus- 
trations of E. funatus indicates considerable 
differences between the two. The latter is more 
tightly coiled and has a thickened arcuate 
columellar lip. It also possesses fewer but 
more strongly developed spiral cords. The 
Lilydale form has less prominent collabral 
growth lines, but these are retrousse over the 
spiral elements, a feature apparently lacking 
in E. funatus. The sutures of O. rotundimur- 
atus are also deeper. 

Comparison of 0. rotundimuratus with Om- 
phalotrochus globosum (Schlotheim) as re- 
described by Lindstrom (1884, p. 160) re- 
veals numerous differences. The former is con- 
siderably lower spired with deeper sutures 
and wider umbilicus. The form from Lilydale 
possesses a straighter columellar lip and less 
rounded outer whorl profile. It also has less 
numerous and weaker spiral sculptural ele- 
ments; nor do these elements exhibit a ser- 
rated to nodose appearance as in 0. globosum. 
The Gotland form also lacks the retrousse 
intersection of the spiral and collabral elements 
as in O. rotundimuratus. 

Superfamily NERITACEA Rafinesque, 1815 
Family Neritopsidae Gray, 1847 

Genus Naticopsis McCoy, 1844 

Subgenus Naticopsis (Naticopsis) McCoy, 

1844 
Type Species: Naticopsis phillipsi McCoy, 
1 844; Lower Carboniferous; Kilcommock, 
Longford, Ireland. 

Range: Lower Devonian to Triassic. The pre- 
sence of a species of this subgenus at Lilydale 
extends the lower limit of its range from 
the Middle Devonian to the Lower Devonian. 

Naticopsis (Naticopsis) lilydalensis 

Cresswell, 1893 

(PI. 1, fig. 9, 10) 
1893 Naticopsis lilydalensis Cresswell, p. 44, pi. 9, 

1913 Craspedostoma lilydalensis (Cresswell): Chap- 
man, p. 227. 

1916 Craspedostoma lilydalensis (Cresswell): Chap- 
man, p. 95, pi. 4, fig. 37. 

Diagnosis: Form of genus with slight spire, 
rounded to only slightly extended base, dis- 



REVISION OF THE GASTROPOD FAUNA 



13 



tinctly auriform aperture, outer lip basal region 
thickened and excavated; slightly arcuate to 
straight, thick columellar lip. 
Description: Medium, low spired, naticiform 
gastropod; whorl profile rounded with upper 
surface slightly flattened; shallow adpressed 
sutures; base rounded to slightly extended; 
without umbilicus; aperture auriform, outer 
lip slightly, irregularly, prosocline; moderately 
thin on upper and outer surfaces but thickens 
considerably on lower surface towards junction 
with the columellar lip, thickened region ex- 
cavated; columellar lip thick, straight to slight- 
ly arcuate; parietal inductura variably devel- 
oped, generally moderately thick; collabral 
lines fine and closely spaced, occasionally irreg- 
ular; infrequently more prominent lines devel- 
oped; no other sculpture. 



Dimensions: 

m 

P948 19 

P949 — 

P37740 30 



Wt 



Hap 



Wap Wh 



22-6 
29 



20 



16 



4 
4 



Location of Types: National Museum of Vic- 
toria. Holotype, P948; Hypotypes P951 and 
P37740. A. W. Cresswell Coll. 
Material: Holotype, 2 hypotypes and 10 other 
specimens. 

Discussion: Absence of the apertural region 
on the holotype has resulted in some confusion 
as to which genus this species should be as- 
signed. Chapman (1913, p. 227 ) ascribed 
this species to the genus Craspedostoma with- 
out reason. Subsequently in 1916, he noted 
(p. 95) that the base had an umbilicus and 
that 'in a supplementary specimen, a part of 
the columellar area of the everted lip is pre- 
served, which shows relationship to the above 
genus'. The hypotype P951 was considered 
by Chapman to possess an umbilicus. This 
specimen is in fact an internal mould, the 
umbilicus being the space that would have 
been occupied by the columella. No specimen 
has been found suggesting the presence of an 
'everted lip'. Chapman also mentioned the 
presence of an obscure cancellated sculpture, 
consisting of flattened spiral and collabral ribs. 
Again no evidence for the presence of spiral 
elements has been found. 



Comparison of N. (N>) lilydalensis with the 
type species reveals that it is slightly higher 
spired but has a less elongate last whorl. The 
form from Lilydale also has a more auriform 
aperture with a straighter columellar lip. The 
lower area of the outer lip is also considerably 
thicker. However, the parietal inductura of 
the type species is thicker. 

Superfamily MURCHISONIACEA Koken, 

1896 
Family Murchisoniidae Koken, 1896 

Genus Murchisonia D'Archiac and De 

Verneuil, 1841 
Subgenus Murchisonia (Murchisonia) 

D'Archiac & De Verneuil, 1841. 
Type Species: Muricites turbinatus Schlotheim, 
1820; Middle Devonian; Stringocephalus lime- 
stone, near Bladbach im Bergischen, Germany. 

Murchisonia (Murchisonia) pritchardi 

(Etheridge), 1898 
(PL 3, fig. 11, 12, 13, 14) 

1898 Goniostropha pritchardi Etheridge, p. 71, pi. 
15, figs. 1-4. 

1913 Murchisonia {Goniostropha) pritchardi Eth- 
eridge; Chapman, p. 227. 

1916 Goniostropha pritchardi Etheridge; Chapman, 
p. 88, pi. 4, fig. 29. 

1916 Cyrtostropha lilydalensis Chapman, p. 87, pi. 4, 
figs. 26-28. 

Diagnosis: Typical form of subgenus but pos- 
sessing spiral sculpture above and below the 
selenizone. 

Description: Medium, high spired, numerous 
whorled gastropod with a selenizone between 
two prominent cords at the angular periphery; 
the whorl face is flat to slightly concave both 
above and below the selenizone; sutures mod- 
erately deep; base rounded; lacking umbilicus; 
columellar lip thin, arcuate and reflexed; junc- 
tion of columellar and other lip not known; 
parietal inductura thin; outer lip with angular 
sinus that forms a slit at the periphery which 
generates the selenizone; from the upper suture 
to the selenizone the outer lip passes posteriorly 
with a moderate obliquity; below the seleni- 
zone it passes forwards to the base less 
strongly; selenizone concave; collabral lines 
fine and weakly developed; sculpture consists 
of a number of spiral cords above and below 
the selenizone; cords more numerous below the 



14 



C B. TASSELL 



selenizone than above it; none of the spiral 
cords are as strong as those bordering the 
selenizone. 



Wt Hap Wap Wh Clu C1I 



8+ 



Dimensions: 

Ht_ 

F.4112 20-1 9-4 7 3 

F.4112 22-5 9-6 56 4-7 7 3 

P935 12-9 7-7 41 3 6 4 3 

P936 16-5 9-5 — — 7 3 10 

P941 18 7 5 5-6 41+ 6-\ — 

P944 31-2 — — — 8+ — — 

P946 230 9-5 5-4 40 8 — — 

Location of Types: 1. Goniostropha pritchardi, 
Australian Museum. Syntypes F.4112. Of 
these 5 specimens, one is that figured by 
Etheridge, as figure 1, plate 15, another is 
that illustrated as figures 2 and 3, plate 15. 
The former of these two specimens is the 
most complete and is here designated the lecto- 
type. National Museum of Victoria. Hypotypes, 
P935-940. A. W. Cresswell Coll. 

2. Cyrtostropha lilydalensis, National Mus- 
eum of Victoria. Holotype, P944. Paratype, 
P946. Hypotypes, P941-3, P945, P947. 
Material: Lectotype, 4 paralectotypes, 13 hypo- 
types and 48 other specimens. 
Discussion: Chapman (1916, p.88) distin- 
guished C. lilydalensis from G. pritchardi on 
the basis of the former's shorter habit, more 
angulate whorls and deeper selenizone. How- 
ever, these features vary independently of each 
other and such variation as does exist is con- 
sidered to represent variation within a popu- 
lation only. 

Chapman (1916, p. 88) suggested that the 
specimen of Murchisonia sp. mentioned by 
Etheridge (1891, p. 129) might be C. lilydal- 
ensis. Re-examination of this specimen indi- 
cates that it is not. Rather it is a poorly pre- 
served and crushed fragment of what appears 
to have been a moderately large specimen of 
Michelia brazieri (Etheridge). 

M. (M.) pritchardi differs from the type 
species noticeably in possessing elements of 
spiral sculpture above and below the seleni- 
zone. The type species has a slightly more 
angular periphery and its growth lines above 
the selenizone pass backwards more obliquely. 
The Lilydale form has a slightly higher peri- 
pheral selenizone. 



Genus Michelia Roemer, 1852 

Type Species: Michelia cylindrica Roemer, 
1854; Devonian; Bockswiese, near Clausthal, 
Germany. 

Discussion: A new Palaeozoic subgenus of the 
Tertiary genus Niso (Risso) was erected by 
Etheridge (1890, p. 62). He considered that 
ultimately it would 'reveal an organization 
differing from Niso in which case I would 
propose for it the name Vetotuba'. 

Although Etheridge (1890, p. 63) mention- 
ed that Niso darwini de Koninck (1876, p. 
127) from Yass had an umbilicus similar to 
N. (Vetotuba) brazieri, no reference to JV. 
darwini was made in the generic description 
and discussion. Thus Knight (1941, p. 382) 
considered Vetotuba brazieri to be the type 
species. 

Knight (1944, p. 459) synonymized Veto- 
tuba with Coelocaulus (Oehlert). This latter 
genus was in turn synonymized by Knight 
et al. (I960, p. 1292) with Michelia, as was 
Vetotuba itself. 

Michelia brazieri (Etheridge), 1890 
(PI. 1, fig. 3, 5, PL 2, fig. 1) 
1890 Niso (Vetotuba') brazieri Etheridge, p. 62, pi. 
8, figs. 4-5, pi. 9, figs. 2-3. 

PLATE 3 

Fig. 1 — Straparollus (Euomphalus ) northi (Eth- 
eridge). P1115, hypotype, X H. Exterior 
of operculum. 

Fig. 2 — Straparollus {Euomphalus) northi ( Eth- 
eridge), P28719, hypotype, X 1£. Basal 
view. 

Fig. 3. — Bellerophon (Bellerophon) cresswelii Eth- 
eridge, P1087, hypotype, X 3. 

Figs. 4-6 — Bellerophon (Bellerophon) cresswelii Eth- 
eridge, F.1327, holotype, X 1 1/3 (approx). 

Fig. 7 — Straparollus (Euomphalus) northi (Eth- 
eridge, P28499, hypotype, X 2/3. Section 
showing transverse partitions developed in 
early whorls. 

Fig. 8 — Straparollus (Euomphalus ) northi (Eth- 
eridge), F.1139e, paratype, X 1 (approx.). 

Fig. 9 — Bellerophon (Bellerophon) cresswelii Eth- 
eridge, P34938, hypotype, X 3. 

Fig. 10 — Gyrodoma etheridgei (Cresswell), P38504, 
hypotype, X 4/5. 

Fig. 11 — Murchisonia (Murchisonia) pritchardi (Eth- 
eridge), F.4112b, paralectotype, X 2i. 

Fig. 12 — Murchisonia (Murchisonia) pritchardi (Eth- 
eridge), F.4112a, lectotype, X 2£. 

Fig. 13 — Murchisonia (Murchisonia) pritchardi 
(Etheridge), F.4112b, paralectotype, X 2£. 

Fig. 14 — Murchisonia (Murchisonia) pritchardi (Eth- 
eridge), P946, hypotype, X2. 



MEM. NAT. MUS. VICT. 37 PLATE 3 








\08> 



«> 




',J»iX 






s 



•i 

5 



m 




***■' 



j* 









■ 

' 6 



N 







REVISION OF THE GASTROPOD FAUNA 



15 



1894 Niso (Vetotuba) brazieri Etheridge; Cresswell, 

p. 158. 
1913 Vetotuba brazieri Etheridge; Chapman, p. 227 

(in part). 
1916 Coelocaulus brazieri (Etheridge); Chapman, p. 

86, pi. 3, figs. 20-22. 
1916 Coelocaulus apicalis Chapman, p. 87, pi. 3, 

figs. 23-24. 
1941 Vetotuba brazieri Etheridge; Knight, p. 382, 

pi. 46, figs. 3a-c. 

Diagnosis: Medium to large, narrowly umbiii- 
cate, cyrtoconoid gastropod, with pseudose- 
lenizone. 

Description: Medium to large, high spired, 
cyrtoconoid gastropod; numerous whorls; 
whorl profile gently convex to nearly flat; 
sutures shallow, impressed; base flatly rounded 
with sub-rounded periphery, narrowly umbili- 
cate; columellar lip straight otherwise inner 
lip unknown; outer lip unknown except that it 
gives rise to a pseudoselenizone that is border- 
ed by two ridges; sculpture known only in 
part; fine collabral lines present on the whorl 
base, umbilicus and lower region of the outer 
whorl surface; the collabral lines swing back- 
wards moderately from the umbilicus and 
continue across the base and onto the lower 
area of the outer whorl surface, where they 
continue backwards a short distance; moder- 
ately thin shell; earlier whorls greatly thick- 
ened by internal secondary deposits. 

Dimensions: All specimens are incomplete. 



Ht 



wt 



Wh 



F.1145 


42 


22 


10 


F.1240 


55 


— 


— 


P1057 


43 


234- 


— 


P1058 


41 


17 


12 


P12842 


52 


20 -f 


— 


P12843 


32 


19+ 


— 


P37755 


53 


20 


10 



Location of Types: 1. Vetotuba brazieri. Aus- 
tralian Museum. Holotype, F.1145 (designated 
by Knight (1941, p.382)). Paratype, F. 1240a. 
National Museum of Victoria. Hypotypes, 
P1057, A. W. Cresswell Coll. P12842, J. S. 
Green Coll. P12843, G. B. Pritchard Coll. 

2. Coelocaulus apicalis, National Museum 
of Victoria. Holotype, P1058. Paratype, P1059. 
A. W. Cresswell Coll. 

Material: Holotype, 1 paratype, 6 hypotypes 
and 52 other specimens. Most of the specimens 
lack their apical region. 



Discussion: Chapman (1916, p. 86) described 
the presence of a 'slit band below median line, 
feebly concave bounded by threads above and 
below'. He considered that PI 2843 exhibited 
the slit band. Re-examination of this specimen 
suggests that the irregular spiral grooves are 
the result of weathering. However, both PI 060 
and P37757 quite clearly possess a pseudosel- 
enizone. This feature is only found on speci- 
mens preserved in the unweathered limestone. 
It is not normally found on specimens that 
have weathered free. 

Chapman (1916, p. 87) erected a new species 
C. apicalis and distinguished it from V. braz- 
ieri on the basis of its smaller size, smaller 
spiral angle, more numerous whorls, particul- 
arly in the apical region, and more regularly 
cylindrical umbilicus. Variation in these char- 
acters in the material from Lilydale is insuffi- 
cient to justify the erection of a new species 
and is no more than that expected in a single 
population. 

Comparison was also made between V. braz- 
ieri and Niso darwini by Chapman (1916, p. 
87). The form from Yass was distinguished 
principally by its smaller size and the more 
slender apical region. Etheridge also noted that 
N. darwini is smaller than the Lilydale form. 
Comparison of the two forms is limited by 
the nature of preservation of the Lilydale speci- 
mens. However, M. brazieri is generally of a 
greater size, particularly length, than the Yass 
form. M. darwini is also more slender in the 
apical region and has a more angular basal 
periphery. 

M. brazieri as it is presently known differs 
principally from the type species in being 
cyrtoconoid in shape. Further comparison be- 
tween the two is limited because of the rel- 
atively poor preservation of both the type 
species and the Lilydale species. One specimen 
P40619 is cryptomphalous, the umbilical reg- 
ion on the base being covered by a swollen 
callus deposit. This feature has only been 
observed in the one specimen. 

Chapman (1907, p. 73) reported M. braz- 
ieri from the limestones of Marble Creek, 
Thomson River, Victoria. However, this 
specimen and material collected more recently 



16 



C. B. TASSELL 



by Talent & Philip (1956, p. 62) is too poorly 
preserved for specific identification. 

Michelia darwini (de Koninck), 1876 
(PL 1. fig. 1, 2) 

1876 Niso darwinii de Koninck, p. 127, pi. 4, figs. 

11, lla-c. 
1898 Vetotuba darwinii (de Koninck); Dunn, p. 

101. English translation of above. 
1916 CoeJocaulus darwinii (de Koninck); Chapman, 

p. 86. 
1941 Niso darwinii de Koninck; Knight, p. 382. 

Diagnosis: Small to medium, narrowly umbili- 
cate cyrtoconoid gastropod. 
Description: Small to medium, high spired, 
cyrtoconoid gastropod; numerous whorls; whorl 
profile gently convex to nearly flat; sutures 
shallow, impressed; base gently rounded with 
angular periphery, narrowly umbilicate; aper- 
ture subrhomboidal; columellar lip straight 
and thin, outer lip thin, straight from the up- 
per suture and passing backwards to the basal 
edge very gently; sculpture unknown; moder- 
ately thin shell. 

Dimensions: All material broken. 

Ht _Wt Wh Hap Wap 



ANU 36851 


210 


11-7 


18 








ANU 36852 


31 5 


13-2 


17 


, 





ANU 36853 


35-5 


18 6 


9 


63 


7-5 


PI 2699 


9-6 


4-4 


7 







PI 2700 


17 4 


5-7 


13 





. . 


P12701 


13 4 


7 6 


6 


— 






Location of Types: 1. Niso darwini, the speci- 
mens figured by de Koninck were destroyed 
by fire when the Garden Palace in Sydney 
was burnt on September 22nd, 1882. 

2. Michelia darwini, Geology Department, 
Australian National University. Hypotypes, 
ANU 36852 and ANU 36853. 
Type Locality: 1. Niso darwini, a black com- 
pact limestone in the Yass District. 

2. Michelia darwini; Chatterton's (1973, 
p. 140) locality B, in the lower half of the 
'Receptaculites' limestone about 400 m east- 
southeast of the homestead on Bloomficld 
Property, Parish of Waroo, near Yass. 
Stratigraphic Range: The 'Receptaculites' Lime- 
stone is considered by Strusz (1972) to be 
Emsian. 

Material: hypotypes and 23 other specimens. 
Discussion: De Koninck (1876, p. 127) gave 



the location of his specimens as a black lime- 
stone in the Yass District. Etheridge (1890, 
p. 63) subsequently gave the location of de 
Koninck's specimens as the Upper Silurian, 
probably Wenlockian beds near Yass. Speci- 
mens from the Shearsby Collection in the 
National Museum of Victoria were collected 
from Portion 208, Parish of Waroo, N.S.W. 
Specimens from the Geology Department, Aus- 
tralian National University are from the local- 
ities B and M of Chatterton (1973, p.140). 
Both these localities are separated from the 
Upper Silurian sediments at Yass by a major 
geological structure. 

The absence of a pseudoselenizone in M. 
darwini is another distinguishing feature be- 
tween the two species. However, the limited 
preservation of this feature in the Lilydale 
material suggests its absence at Yass may be 
due to the nature of preservation. 

Family Plethospiridae Wenz, 1938 

Genus Gyrodoma Etheridge, 1898 

Type Species: Eunema etheridgei Cresswell, 

1893; Lower Devonian; Lilydale Limestone, 

Lilydale. 

LOCATION OF TYPES 
TYPE LOCALITY 

Gyrodoma etheridgei (Cresswell), 1893 

(PL 2, fig. 2, 9. PL 3, fig. 10) 

1893 Eunema etheridgei Cresswell, p. 42, pi. 8, fig. 2. 
1898 Gyrodoma etheridgei (Cresswell); Etheridge, 

p. 72, pi. 16, fig. 1. 
1913 Gxrodoma etheridgei (Cresswell); Chapman, 

p. 227. 
1941 Gyrodoma etheridgei (Cresswell); Knight, p. 

138, pi. 4, figs. la-c. 
1960 1 Gxrodoma etheridgei (Cresswell); Knight et 

at., p.I296, fig. 192, 2. 

Diagnosis: Large, high-spired gastropod with 
rounded whorls and deep sutures; selenizone 
broad, flat; sculpture, numerous spiral threads; 
typically threads also on selenizone. 
Description: Large, high spired gastropod; 
whorl profile well rounded with deeply im- 
pressed sutures; peripheral selenizone at mid- 
whorl; minutely umbilicate; inner lip concave; 
columellar lip thickened and reflexed extends 
as inductura to envelope the umbilicus; pariet- 
al inductura thin and extensively developed; 
outer lip not known in detail, no growth lines 



REVISION OF THE GASTROPOD FAUNA 



17 



known; outer lip gives rise to a broad, flat, 
sculptured selenizone, bounded by threads 
slightly more prominent than those constituting 
the remaining whorl sculpture; the selenizone 
in the larger whorls generally has a single 
fine median spiral thread dividing it into two 
equal parts; the selenizone in the earlier whorls 
may have up to three threads so dividing it; 
sculpture composed of numerous fine spiral 
threads of two or more orders above and 
below the selenizone; more threads below the 
selenizone than above it; threads continue into 
the umbilicus. 



Dimensions: 
Ht 



Wt 



P10187 
P38503 
P38504 
F.2542 



Hap 



Wap Wh 



29+ 
62 
51 
62 



35 + 
31 
25 
29 



— 5+ 

— 4+ 

— 5 + 



Location of Types: National Museum of Vic- 
toria. Holotype, P10187. Presented by the 
Rev. A. W. Cresswell to the National Museum 
of Victoria on September 9, 1908. Hypotype 
P38504. Australian Museum. Hypotype, F.25 
42. 

Material: Holotype, 2 hypotypes and 24 other 
specimens. 

Discussion: As figured by Cresswell the holo- 
type has vertical growth lines on the divided 
selenizone. However, on examination of this 
specimen, no suggestion of their presence was 
found. Etheridge noted that while the holotype 
has a divided selenizone, hypotype F.2542 is 
without. Typically the selenizone has one or 
more threads developed on it. 

Knight (1941, p. 138) designated as the 
holotype the left hand illustration in fig. 2, PI. 
8 of Cresswell (1893). However, it is con- 
sidered both illustrations are of the same speci- 
men. This is certainly true for the other 
specimens on the plate Tremanotus pritchardi 
and Siluriphorus antiquus which are similarly 
figured. Comparison of the holotype with the 
right-hand illustration suggests that the smaller 
whorl of the holotype has been broken sub- 
sequently. Supporting this is the presence of 
a fresh area of matrix and shell on the upper 
surface of the holotype. 



Superfamily PSEUDOPHORACEA S. A. 

Miller, 1889 
Family Pseudophoridae S. A. Miller, 

1889 
Genus Scalaetrochus Etheridge, 1890 

Type Species: Trochus (Scalaetrochus) lin- 
strotni Etheridge, 1890; Lower Devonian; 
Lilydale Limestone, Lilydale. 

Scalaetrochus lindstromi Etheridge, 1890 
(PI. 1, fig. 13, 15, 16, 18. PL 2, fig. 5) 

1890 Trochus (Scalaetrochus) lindstromi Etheridge, 

p. 66, pi. 8, figs. 1-2. 
1913 Trochus (Scalaetrochus) Undstroemi Etheridge, 

Chapman, p. 228. 
1916 Scalaetrochus Undstroemi Etheridge; Chapman, 

p. 94. 
1941 Scalaetrochus lindstromi Etheridge; Knight, p. 

306, pi. 59, figs. 3a-d. 

1959 Scalaetrochus lindstromi Etheridge; Philip and 
Talent, p. 53, pi. 8, figs. 3-8. 

1960 Scalaetrochus Undstroemi Etheridge; Knight 
et al., p. 1298, fig. 195, 1. 

Diagnosis: Large trochiform low whorled gas- 
tropod with mildly concave cryptomphalous 
base and narrow peripheral frill; callus deposit 
beginning in aperture and filling peripheral 
angle; deposit variably developed in umbilicus, 
collabral lines moderately prosocline on outer 
whorl surfaces. 

Description: Large trochiform cryptomphalous 
gastropod with mildly concave base; irregular 
sutures flush to slightly protruding; whorl 
profile gently to moderately concave; periphery 
angular, forming narrow frill; columellar lip 
thickened and strongly oblique outwards; 
parietal inductura thin or wanting; thickened 
outer lip moderately prosocline from the upper 
suture to the basal periphery, it continues 
obliquely across the base to the columellar 
Up; the columellar and outer lips on the base 
are strongly concave; a callus is variably devel- 
oped in the umbilicus, ranging from near 
absence in the umbilicus to complete infilling 
of the umbilicus; umbilical callus continuous 
with the callus which occupies the peripheral 
angle; this material is deposited anterior to 
the aperture; collabral, growth lines, fine to 
slightly foliaceous on the outer whorl surface; 
collabral lines on the base are fine and when 
the umbilicus is open continue into it; occas- 
ional rugae occur on the base. 



18 



C. B. TASSELL 



Dimensions: 



Ht 



Wt 



Hap 



Wap Wh 



F.1137 

P294 
P921 
P16024 
P38505 



47 72 18 36 5 

40 64 — 25+ 6 

_ 60 — — — 

43 58 — — 6 

39 64 — — 5 

P38506 36+ 64 15 30 5 

Location of Types: Australian Museum. Holo- 
type, F.1137. National Museum of Victoria. 
Hypotypes, P38505 and P39279. E. D. Gill 
Coll. 

Material: Holotype, 2 hypotypes and 37 other 
specimens. 

Discussion: As originally described by Ether- 
idge, S. lindstromi was without an umbilicus. 
Philip & Talent (1959, p. 53) indicated that 
it is cryptomphalous. However, the degree of 
development of the callus is more variable 
than that intimated by Chapman (1916, p. 
93) or stated by Philip and Talent. 

Specimens have frequently been crushed, 
the base suffering the greatest distortion and 
damage. This occurs generally where the 
whorl is thinnest, at the inner edge of the 
peripheral angle thickening. 

Genus Siluriphorus Cossmann, 1918 
Type Species: Trochus gotlandicus Lindstrom, 
1884; Middle Silurian; the canal near Wes- 
toos in Hall, Gotland, Sweden. 
Range: Middle Silurian to Lower Devonian. 
The presence of a species of this genus at 
Lilydale extends the upper limit of its range 
from the Middle Silurian to Lower Devonian. 
Distribution: Europe and Australia. The pre- 
sence at Lilydale of a species of this genus 
extends the generic range to include Australia. 
Discussion: Philip and Talent (1959, p. 53) 
synonymized Siluriphorus with Scalaetrochus 
Etheridge. They considered the differences in 
size, deflection of the outer lip and sculpture 
between the type species are specific rather 
than generic. 

Close examination of the type species sug- 
gests that until there is a review of the entire 
family both genera are valid. 5. lindstromi 
can be distinguished from S. gotlandicus by 
the callus deposit which is not only present 
in the umbilicus but also extends to fill the 
peripheral angle in advance of the aperture. 



The periphery of the Lilydale form is angular 
and extends to form a narrow frill, whereas 
the Gotland form is characterised by a more 
variable periphery. The holotype of Siluripho- 
rus gotlandicus has a blunt angular periphery 
but in some specimens a blunt frill-like border 
is developed. Similarly the collabral lines on 
the upper whorl surface of S. gotlandicus are 
more variable, the holotype having quite 
coarse, strong, irregular imbricating lamellae. 
S. lindstromi has very much finer more regular, 
foliaceous collabral lines. These lines are also 
more arched and are not directed backwards 
as obliquely as in the Gotland form. 5. gotland- 
icus is characterized by an almost flat to gently 
convex whorl profile between obscure shallow 
sutures, whereas S. lindstromi has more prom- 
inent irregular, flush to slightly protruding 
sutures and a distinctly concave whorl profile. 

Siluriphorus antiquus (Cresswell), 1893 
(PI. 2, fig. 4, 6) 

1893 Stomatia antiqua Cresswell, p. 43, pi. 8, fig. 3. 

1894 gen. indet. Cresswell, p. 157. 

1913 Trochus {Scalaetrochus) antiquus (Cress- 
well); Chapman, p. 228. 

1916 Scalaetrochus antiquus (Cresswell); Chapman, 
p. 93. 

Diagnosis: Typical form of genus with sub- 
rounded periphery and fine, regular, imbri- 
cating growth lamellae. 

Description: Medium, trochiform gastropod 
with few low whorls and weakly impressed 
sutures; whorl profile gently convex to nearly 
flat between sutures; basal periphery sub- 
angular; base flat to slightly rounded, details 
of umbilicus not known but probably cryptom- 
phalous; aperture probably avoidal; columellar 
lip thickened and strongly oblique outwards; 
parietal inductura thin; outer lip strongly ob- 
lique backwards from the upper suture to the 
periphery, only very weakly prosocline; ob- 
liquity maintained across the periphery onto 
the whorl base; numerous fine imbricate col- 
labral lamellae on outer whorl surface; no 
thickening of the peripheral region. 
Dimensions: 



Ht 



Wt 



Hap Wap 



W 



P917 


14+ 


39 — 


P916 


36+ 


44+ — 


P918 


13 + 


36 — 



2+ 
3+ 
2+ 



REVISION OF THE GASTROPOD FAUNA 



19 



Location of Types: National Museum of Vic- 
toria. Holotype, P917. Hypotype, P918. A. 
W. Cresswell Coll 

Material: Holotype, hypotype and one other 
specimen. All the specimens are incomplete 
and crushed to varying degrees. 
Discussion: The limited number of specimens 
and their crushed state precludes a complete 
description of the species. 

The type species exhibits considerable vari- 
ation in the strength and coarseness of the 
growth lamellae, Lindstrom (1884, pi. 14, figs. 
1-11). Knight (1941, p. 318) noted that the 
holotype is a specimen with coarse, strong 
imbricating lamellae. Although cryptomphalous 
there is also considerable variation in the form 
of the umbilical callus. 

S. antiquus in comparison with the holotype 
of the type species has weaker, more regular 
imbricating growth lamellae on the outer whorl 
surface. The type species as exemplified by the 
holotype has a more angular periphery than 
the Lilydale form. None of the specimens from 
Lilydale possesses a bourrelet as found in 
some of the specimens from Gotland. 

Superfamily LOXONEMATACEA Koken, 

1889. 

Family Loxonematidae Koken, 1889. 

Genus Loxonema Phillips, 1841. 

Type Species: Terebra? sinuosa J. deC. Sower- 

by, 1839; Middle Silurian; Garden House, 

near Aymestry, Shropshire, Britain. 

Loxonema australis (Chapman), 1916 
(PI. 1, fig. 4, PL 2, fig. 8) 

1916 Loxonema sinuosa var. australis Chapman, p. 

96, pi. 5, fig. 39. 
1949 Loxonema australis (Chapman); Gill, p. 111. 
Diagnosis: Medium, intermediate form be- 
tween Loxonema and Palaeozygopleura. As 
a member of the former genus it possesses a 
shallow sinus which deepens slightly with age 
and sutures of moderate depth. 
Description: Medium, high spired numerous 
whorled gastropod with a shallow rounded 
sinus in the outer lip; whorl profile gently 
arched, periphery at mid-whorl; sutures mod- 
erately impressed; base rounded; lacking um- 
bilicus; inner lip concave; columellar lip short, 



thickened and reflexed; parietal inductura thin; 
outer lip with a very shallow wide sinus devel- 
oped slightly above mid-whorl; from the upper 
suture the growth lines are very steeply pro- 
socline to the middle of the sinus, before pass- 
ing more gently forwards to the edge of the 
base where they curve prosocyrtly to the 
centre; sinus deepens slightly with growth; out- 
er lip of moderate thickness, except towards 
the junction with the columellar lip where 
thicker; collabral sculpture consisting of costae 
and costellae; protoconch unknown. 
Dimensions: 



Ht 



Wt 



Hap 



Wap 



Wh 



P12851 


22 


19 


P38507 


47+ 


19 


P3850S 


31 + 


14 



8 



12 



1 

5+ 
5+ 



Location of Types: National Museum of Vic- 
toria. Holotype, P12851, J. S. Green Coll. 
Hypotype, P38508. 

Material: Holotype, hypotype and 8 other 
specimens. 

Discussion: Comparison of L. australis with 
the type species L. sinuosa reveals that the form 
from Lilydale has a distinctly shallower and 
wider labral sinus. It also has a coarser col- 
labral sculpture which weakens slightly on the 
base and is slightly larger. There is no evidence 
for 'a tendency to form a faint nodose shelf 
near the basal part of the whorl', the only 
distinguishing feature between the two species 
noted by Chapman. 

The type species of Palaeozygopleura, P. 
alinae Perner, when compared with the Lily- 
dale form has a noticeably shallower and 
wider labral sinus. It also is considerably 
smaller and has much shallower sutures than 
L. australis. Thus the form from Lilydale oc- 
cupies an intermediate position between these 
two genera as regards the depth of the labral 
sinus. Such a situation is not unexpected. 
Knight (1930) , postulated that Loxonema 
gave rise to the Carboniferous Pseudozy- 
gopleuridae through an unknown Pseu- 
dozygopleura-Yike form that existed in the 
Devonian or Lower Carboniferous. This fonn 
was most probably represented by the Pala- 
eozygopleuridae and more particularly 
Palaeozygopleura. Thein and Nitecki (1974, 



20 



C. B. TASSELL 



p. 29) considered 'that the Palaeozygopleuri- 
dae evolved from Loxonema directly during 
the Devonian or Mississippian and gave rise to 
the Pseudozygoplcuridae that flourished in the 
Pennsylvanian and Permian'. 

The Mississippian form L. knighti Yochelson 
(1962, pi. 17, fig. 11) possesses in its inter- 
mediate growth a shallow wide labral sinus 
and moderately strong collabral sculpture com- 
parable to the intermediate growth stage in 
L. auslralis. In its mature form the American 
species possesses the characteristically deep 
labral sinus of the genus. However, the labral 
sinus of the mature Lilydale form does not 
deepen as much, although it is deeper than 
that of the immature American form. The 
species are comparable in size. 

Two other genera within the family Loxon- 
ematidae also possess shallow sutures, they 
being Stylonema and Aulacostrepsis. Both 
occur in the Lower Devonian. However, Sty- 
lonema is characterized by a very slender shell 
and the similar Aulacostrepsis also has a very 
small umbilicus. 

Although the Lilydale form is obviously in- 
termediate between the two genera Loxonema 
and Paiaeozygopleura it is for the present as- 
signed to the former genus because of its 
moderately deep labral sinus, which deepens 
slightly with growth, size, moderate sutures 
and the nature of the labral sinus in other 
genera of the family Loxonematidae. 

Gill (1949, p. Ill) in redescribing the holo- 
type considered that Chapman s variety was 
in fact a new species distinguished by the 
coarser nature of the sculpture and the 
straighter costellae. He also assigned the form 
from Sandy's Creek, Parish of Nungatta, Gipps- 
land to this species noting that it is smaller 
and that the ornamentation is proportionally 
finer. However, Talent (1963, p. 102) con- 
sidered that the form from Sandy's Creek 
differs sufficiently from L. australis to be con- 
sidered a new species, this species being dis- 
tinguished by its overall size. Never exceeding 
10 mm in width, it is much smaller than L. 
australis. The whorls in the form from Sandy's 
Creek are also proportionally higher than 
those of L. australis. This unnamed species 



of Loxonema is also found in the mudstones 
at Loyola. 

References 

Baths, D. E., 1972. A new Devonian crinoid from 

Australia. Palaeontology 15 (2): 326-335. 
Boucot, A. J., 1975. Evolution and Extinction Rate 

Controls. (Developments in Palaeontology and 

Stratigraphy, 1.) Elsevier. 
Chapman, F., 1907. Newer Silurian fossils of eastern 

Victoria, Part 1 Rec. geol. Surv. Vict. 2 (I); 

67-80. 
, 1913. On the Palaeontology of the Silurian 

of Victoria. Rep. Australas. Ass. Advmt. Sci. 14: 

207-235. 

1916. New or little-known Victorian fossils 



in the National Museum. Proc. R. Soc. Vict. 29: 

75-103. 
Chatterton, B. D. E., 1973. Brachiopods of the 

Murrumbidgee Group, Taemas, New South 

Wales. Bull. Bur. Miner. Resour. Geol. Geophys. 

Aust. 137. 
Clarke, J. M. and Rudemann, R. 1903. Guclph 

fauna in the State of New York. Mem. N.Y. St. 

Mus. nat. Hist. 5. 
Cresswell, A. W., 1885. In Anonymous, The 

Queens Birthday excursion to Lilydale. Victorian 

Nat. 2 (3): 33-36. 
, 1893. Notes on the Lilydale Limestone. 

Proc. R. Soc. Vict. 5: 38-44. 

-, 1894. Additional notes on the Lilydale 



Limestone. Proc. R. Soc. Vict. 6: 156-159. 

Etheridge, R., Jr., 1890. Descriptions of Upper Silu- 
rian fossils from the Lilydale Limestone, Upper 
Yarra District, Victoria. Rec* Aust. Mus. 1 (3): 
60-67. 

, 1891. Further descriptions of the Upper 

Silurian fossils from the Lilydale Limestone, 
Upper Yarra District, Victoria. Rec. Aust. Mus. 
1 (7): 125-130. 

- 1894. An operculum from the Lilydale 



Limestone. Proc. R. Soc. Vict. 6: 150-166. 

- 1898. New or little-known Lower Palaeo- 



zoic Gastropoda in the collection of the Aus- 
tralian Museum. Rec. Aust. Mus. 3 (4): 71-77. 
Gill, E. D., 1949. Devonian fossils from Sandy's 

Creek, Gippsland, Victoria. Man. nam. Mus. Vic. 

16: 91-115. 
Horny, R. L, 1962. New Genera of Bohemian Lower 

Paleozoic Bellerophontina. Vest, listred. Ust. 

geol. 37 (6): 473-476. 
, 1963. Lower Paleozoic Bellerophontina 

(Gastropoda) of Bohemia. Sb. geol. Ved. Paleon- 

tologie 2: 57-164. 
Jhaveri, R. B., 1969. Unterdevonische Gastropoden 

aus den Karnischen Alpen. Palaeontographica 133 

(A): 146-176. 
Knight, J. B., 1930. The gastropods of the St. Louis, 

Missouri, Pennsylvanian outlier: the Pseudozy- 

goplcurinae. /. Paleont. 4 suppl. 1. 
j 1941. Paleozoic Gastropod Genotypes. 

Spec. pap. geol. Soc. Am. 32. 

-, 1944. In Shimer, H. W. and Shrock, R. R., 



Index fossils of North America. New York, John 
Wiley & Sons. 



REVISION OF THE GASTROPOD FAUNA 



21 



Knight, J. B., Batten, R. L., and Yochelson, E. L., 
1960. Descriptions of Palaeozoic Gastropoda. In 
Moore, R. C, ed. Treatise on invertebrate paleon- 
tology: I. Mollusca (1), Univer. Kansas Press. 

Koninck, L. G. de, 1876. Recherches sur les Fossiles 
Paleozoiques de la Nouvelle-Galles du Sud (Aus- 
tralie). Mem. Soc. Roy. Set. Liege 2, 6. (Trans- 
lated, 1898, as: Descriptions of the Palaeozoic 
fossils of New South Wales (Australia)). Mem. 
geol. Surv. N.S.W., Palaeont. 6. 

Lindstrom, G., 1884. On the Silurian Gastropoda 
and Pteropoda of Gotland. K. sevenska Vetensk- 
Akad. Hand I. 19 (6). 

Linsley, R. M., 1968. Gastropods of the Middle 
Devonian Anderdon Limestone. Bull. Am. 
Paleont. 54 (244). 

Manten, A. A., 1971. Silurian Reefs of Gotland. 
(Developments in Sedimentology, 13.) Elsevier. 

Oehlert, D. P., 1888. Descriptions de quelques 
especes devoniennes du department de la May- 
enne. Bull. Soc. Etud. sclent. Angers. 1887, 
65-120. 

Philip, G. M., 1974. Biostratigraphic procedures and 
correlation in the Tasman geosynclinal zone. In 
Denmead, A. K., Tweedale, G. W., Wilson, 
A. F. (Eds). The Tasman Geosyncline — a sym- 
posium. Geol. Soc. Aust. Qld. Div. 

Philip, G. M. and Talent, J. A., 1959. The gas- 
tropod genera Liomphalus Chapman and Scalae- 
trochus Etheridge. J. Paleont. 33: 50-54. 

Schlotheim, E. F. von., 1820. Die Petrefactenkunde 
auf ihrem jetzigen standpunkte durch die Besch- 
reibung seiner Sammlung versteinerter und 



fossiler Uberreste des Thier-und Pflanzenreichs 

der Vorwelt erldutert. Gotha. 
Sowerby, J. DeC, 1823. The mineral conchology 

of Great Britain. London, 5: 1-168. 
Spitz, A., 1907. Die Gastropoden des Karnischen 

unterdevon. Beitr. Paldont. Geol. Ost-Ung. 20: 

115-190. 
Strusz, D. L„ 1972. Correlation of the Lower De- 
vonian Rocks of Australasia. /. geol. Soc. Aust. 

18 (4): 427-455. 
Talent, J. A., 1963. The Devonian of the Mitchell 

and Wentworth Rivers. Mem. geol. Surv. Vict. 

23. 
, and Philip, G. M., 1956. Siluro-Devonian 

Mollusca from Marble Creek, Thomson River, 

Victoria. Proc. R. Soc. Vict. 68: 57-71. 
Thein, M. L. and Nitecki, M. H., 1974. Chesterian 

(Upper Mississippian) Gastropoda of the Illinois 

Basin. Fieldiana, Geol. 34. 
Thompson, E. H., 1970. Morphology and taxonomy 

of Cvclonema Hall (Gastropoda), Bull. Am. 

Paleont. 58 (261). 
Yochelson, E. L., 1962. Gastropods from the Red- 
wall Limestone (Mississippian) in Arizona. /. 

Paleont. 36: 74-80. 
, and Dutro, J. T. 1960. Late Paleozoic 

Gastropoda from Northern Alaska. Prof. Pap. 

U.S. geol. Surv. 334-D. 

-, and Linsley, R. M., 1972. Opercula of 



two gastropods from the Lilydale Limestone 
(Early Devonian) of Victoria, Australia. Mem. 
natn. Mus. Vic. 33: 1-13. 



A NEW SPECIES OF HEMIERGIS (SCINCIDAE: LYGOSOMINAE) FROM 

VICTORIA 

By A. J. Coventry 

Officer-in-Charge, Herpetology, National Museum of Victoria 

Abstract 

A new species of lygosomid skink {Hemiergis millewae) is described, from the Victorian 
Mallee Region. 



Introduction 

Field work in the Victorian Mallee since 
1973, has revealed the presence of a small, 
pentadactyl, lygosomid skink, not previously 
known from this State. The species fits into 
the genus Hemiergis Wagler, 1830, as recog- 
nized by Greer (1967), but does not fit the 
only pentadactyl species (//. initiale Werner, 
1910). A thorough search of the National 
Musuem of Victoria collections brought to light 
a further five specimens, which had previously 
been identified as Anotis maccoyi (Lucas and 
Frost). 

The genus Hemiergis was first erected by 
Wagler in 1830,and has since been redefined by 
many authors, principally Gray (1845), Bou- 
lenger (1887), Mittleman (1952), Greer 
(1967), Cogger (1975) and Storr (in Press). 
Although Cogger and Storr provide the most 
recent definitions, Greer's concept of the genus 
is accepted here. 

Gray provided the first detailed diagnosis of 
the genus, restricting it to skinks with trans- 
parent lower eyelids, and digits 3-3. He also 
considered the presence of paired frontoparie- 
tals to be diagnostic, a character which Storr 
has shown to be variable within the genus, 
even at the specific level. Mittleman restricted 
the genus to species having 4-4 or less digits, 
thus excluding Werner's 1910 pentadactyl spe- 
cies H. initiale. Greer's diagnosis accepted #. 
initiale as a true Hemiergis, thus expanding 
the genus to include pentadactyl species. Cog- 
ger's definition contradicted Greer's in that it 
included . . . 'ear opening usually absent, its 
position is usually indicated by a slight depres- 
sion (a minute opening in one species)'. All 
other workers have agreed that the ear open- 
ing is in fact covered by scales. Cogger's move 



was made to allow the inclusion of Siaphos 
maccoyi Lucas and Frost into Hemiergis, thus 
making Hemiergis polyphyletic, as Cogger 
recognized when he said . . . 'the genus as 
recognized here is almost certainly composite, 
but relationships are obscure'. 

Storr's definition basically agrees with that 
of Greer, differing in that Storr said that all 
species have a complete series of suboculars, 
whereas Greer stated that the subocular row 
was 'complete, except in H. initiale'. In addi- 
tion to the above characters, Greer also sepa- 
rated Hemiergis from Lerista by the presence 
of four supraoculars as against 2-4, usually 3, 
paired rather than single supradigital scales 
on the fourth toe, and non-enlarged as against 
greatly enlarged nasal scales. 

Genus Hemiergis Wagler, 1830 

Hemiergis Wagler, 1830, Nat. syst. amphib.. p. 160. 
Type-species Zygnis decresiensis Fitzinger, 1826. 

Diagnosis: Small elongate, short-limbed skinks. 

Lower eyelid movable, with a transparent disc. 

Digits 5-5 to 2-2, normally equal numbers on 

fore and hind limbs. Subdigital lamellae less 

than 16. Subocular row complete in all non- 

pentadactyl species. Ear aperture absent, ear 

indicated by a depression. Supradigital scales 

paired. 

Hemiergis millewae sp. nov. 
Fig. 1 

Holotype: D47410, adult male in the National Mu- 
seum of Victoria, collected at Millewa South bore, 
Vic., in 34°56'28"S: 141°4'E on 15.XI.1975 by 
A. J. Coventry and P. Mather. 

Description: Snout- Vent (S.V.) length 43-5 

mm. Length of tail (intact) 75-3 mm, 173% 

of S.-V. length. Total length 1188 mm. Length 

of hind limb 130 mm, 30% of S.-V. length. 

Length of fourth toe 4-7 mm, 36% of hind 



23 



24 



A. J. COVENTRY 



limb length. No supra or postnasal scales. 
Rostral and frontonasal in fairly broad contact. 
Frontal and frontonasal in narrow contact. Pre- 
frontals large, just failing to meet: contacting 
the frontonasal, anterior and posterior loreals, 
first supraciliary and frontal. Two loreals, 
large and subequal. Frontoparietal entire, in- 
terparietal separate, large, almost half the size 
of the frontoparietal. Parietals large, barely 
contact along the midline. One pair of enlarged 
nuchals, followed by a second, single nuchal 
on the left hand side. Three enlarged temporals, 
the upper largest. Four supraoculars, the 
second the largest. Seven supraciliaries, seven 
upper ciliaries, the third to fifth largest. Nine 
lower ciliaries. Lower eyelid movable with an 
extremely large transparent palpebral disc bor- 
dered above by the lower ciliaries but otherwise 
surrounded by small granular scales. 

Length of eye 1-6 mm, length of disc 11 
mm, 69% of eye length. Seven upper labials, 
the fifth subocular and completely interrupting 
the subocular series, of which two are anterior 
to, and three posterior to the fifth upper labial. 
Seven lower ciliaries. Ear opening completely 
covered by scales, indicated by a depression. A 
pair of enlarged preanal scales. Limbs short, 
pendactyl, when adpressed, failing to meet by 
approximately 25% of the distance between 
the axilla and the groin. Subdigital lamellae 
dark coloured, undivided and smooth, 12 under 
the fourth toe. Midbody scales smooth, in 22 
rows. 

Colour in Life; Uniform dark olive brown dor- 
sally, with no trace of spots, striations or lines. 
A burnt orange dorso-lateral stripe, approxi- 
mately two scales wide, commencing above and 
behind the ear, and extending to the hind 
limbs. Lateral surfaces off-white, and ventral 
surfaces pale yellow. Dorsal and lateral sur- 
faces of tail similar to mid-dorsal colour, ventral 
surfaces of tail off-white, with darker spots. 
Chin whitish, each scale bordered by dark 
brown. 

Colour in Alcohol: Drab brown dorsally, 
whitish laterally, and light grey ventrally. There 
is almost no trace of the dorso-lateral stripe. 
Paratypes: Fifteen specimens in the National 
Museum of Victoria as follows: 




1cm 




1cm 



Fig. 1 — Head shields of holotype of Hemiergis mille- 
wae D47410, dorsal and lateral views. 

D33348 sex undetermined, D33349 male, 0-3 km S 
of Millewa South Bore, collected G. Barnes 
31.8.1973; the remaining thirteen specimens, all 
from Millewa South Bore as follows: 

D33341 — 2 sex undetermined, collected by G. 
Barnes 29.8.1973; 

D38847 sex undetermined, collected A. J. Coventry 
4.4.1974. 

D40169 sex undetermined, collected A. J. Coventry 
24.9.1974; 

D47394— 6 males, D47397 female, collected A. J. 
Coventry and P. Mather 13.11.1975; 

D47398 — 400 females, collected A. J. Coventry and 
P. Mather 14.11.1975; 

D47418 male, and D47419 female, collected A. J. 
Coventry and P. Mather 16.11.1975. 

Description of Paratypes; As for holotype ex- 
cepting as follows: S.-V. lengths 41*7-58-6 
mm (mean 502); hind limb length 11 1-13-7 
mm (mean 12-3); percentage of S.-V. length 
231-281 (mean 24-7); length of 4th toe 
4-5-5-4 mm (mean 4-8) percentage of hind 
limb length 35-7-420 (mean 390). Upper 



NEW SPECIES OF HEMIERGIS 



25 



ciliaries 8-10 (mean 91), lamellae under 4th 
toe 12-14 mean (12-8). 

Nuchals: one pair (4 specimens), two pairs 
(2 specimens), two pairs plus a third on right 
side (1 specimen), two pairs plus a third on 
left side (1 specimen). D33348 has a damaged 
first toe on the left hind foot; D40169 has the 
right fore foot missing; D47395 has the second 
and third fingers of the right fore limb damaged; 
D47397 has a damaged left fore foot, the 
second and third fingers, which arise from a 
common base, being truncated to appear as a 
single digit; D47398 has the third toe on the 
right hind foot truncated; D47418 has a dam- 
aged fourth toe on the left hind limb. 

Colour of all these specimens, in alcohol, 
similar to the holotype, excepting that in many 
of them, the dorso-lateral stripe is completely 
missing. 

OTHER SPECIMENS EXAMINED 
D1552-3, D1556 from Purnong, S.A., 
Dl 1767-8 from Nonning, S.A., and D47409, 
a juvenile, same data as for holotype. 
Ecology; Little is known of the ecology of this 
species. It resides in porcupine grass {Triodia 
sp.) in sandy soil supporting a fairly heavy 
cover of mallee scrub. It never appears to 
emerge from the Triodia, and was only located 
either by burning or the ripping out of this 
grass. No activity was observed at any time 



of the day or night, although other species of 
reptiles (e.g. Ctenotus brachyonyx, Menetia 
greyi, Amphibolous barbatus, Amphibolous 
fordi, Delma inornata and Lialis burtoni) 
were active during the time spent collecting the 
type series. Obviously //. millewae is a thig- 
motherm, dependent upon Triodia, which one 
assumes supplies its food source in the many 
forms of invertebrates co-habiting with it. 



Key to Hemiergis millewae and other southern 
short limbed skinks 

1. Ear opening visible, not covered 

by scales 2 

Ear covered by scales, indicated by 

a depression 3 

2. Nasals enlarged, meet or almost meet be- 
hind rostral Lerista species 

Nasals not enlarged Anotis maccoyi 

3. Digits 5-5 4 

Digits 4-4 or less .... all other known 

Hemiergis 

4. Hind limb at least 20% of S.-V. length, 

subcaudals 12 or more H. millewae 

Hind limb less than 20% of S.-V. length, 
subcaudals 11 or less H. initiale 

This species is named in recognition of the 
locality where the type series was collected. 



TABLE 1 

Showing comparative lengths of hind limbs, and lamellae under longest toe of 

H. millewae and its allies 



SPECIES 


N 


Hind Limb % S.-V. length 


Lamellae under longest 
toe 




Max. 


Min. 


Mean 


No. 


Mean 


H. millewae 
H. initiale 
H. peroni 
H. decresiensis 
A. maccoyi 


16 
21 
18 
24 
21 


29-9 
18-7 
23-6 
16-5 
19-5 


210 
13-2 
17-7 
111 
151 


25-1 
160 
20-2 
13-8 
17-2 


12-14 
7-10 

10-14 
7-8 
8-9 


12-8 
8-8 

10-7 
7-0 
8-3 



26 



A. J. COVENTRY 



Addendum 

Through the courtesy of Dr. T. F. Houston 
of the South Australian Museum, I have been 
able to examine an additional 119 South Aus- 
tralian specimens of this species, which are 
under his care. These are as follows: — 
R112 Purnong; R3044 A — F near Siam 
Station woolshed, 19.3.1950 under Triodia; 
R3069 A — Z Birthday Well, Cariewerloo 
Station 11.3.1950; R3855 & R1076O — 1 24 
km N. Poochera 15.6.1956; R3860, R10767 
— 73 & R10775 — 6 Kondoolka Turnoff, 
Gawler Ranges 17.6.1956; R5377 & R10762 
6 Gawler Ranges March 1963; R10733 — 
58 Mamblyn 29.4.1969; R11284 21 km N. E. 
Blanchetown 16.2.1969 in sandy soil under a 
spinifex bush in mallee scrub; R 12490 A — 
Z Miccollo Hill, Siam Station 32° 32' S.: 136° 
36' E. 20.4.1971 — ex Triodia Bush; R12619 
8 km from Bakara 34° 4' S.: 139° 45' E. 
January 1970; R13012 A — G 13 km S 
Alawoona36° 6' S.: 140° 32' E. 24.1.1972 — 
ex Triodia bushes; R13099 A — C near Oul- 
nina homestead, Olary Ridge, 32° 34' S.: 139° 
53' E. March — April 1972; R13381 Hiltaba 
Reservoir 32° 10' S.: 135° 4' E. 26.8.1972; 
R13704 1.5 km S. of Mulgathing Rocks 
30° 7' S.: 134° 0' E. 23.4.1973 in Granite 
outcrop; R14766 10 km E., 7.5 km N. of 
Blanchetown 28.8.1975 low dune mallee. 

Of these specimens R3069B, R11284 & 
R13012B each have one foetus in utero 
R3069M, R13012A & D each have two foe- 



tuses in utero, and R3069X, R10736 & R107 
48 are hatchlings. This confirms that the spec- 
ies is viviparous, having one or two young per 
litter, the young being born in late summer or 
early autumn. In these series the midbody scale 
rows vary from 22 — 24. 

Acknowledgements 

The author wishes to thank Dr G. M. Storr 
of the Western Australian Museum for the 
loan of comparative material of Hemiergis 
initiate, and for permission to quote from his 
unpublished manuscript. Mr P. A. Rawlinson, 
Latrobe University for help, encouragement 
and critical reading of the manuscript. Mr P. 
Mather assisted in both field and laboratory, 
and Miss L. Leatham assisted in the laboratory. 
Miss R. J. Plant for the drawings of head scales. 

Bibliography 

Boulenger, G. A., 1887. Catalogue of Lizards in the 
British Museum. London. 

Cogger, H. G., 1975. Reptiles and Amphibians of 
Australia. Reed, Sydney. 

Gray, J. E., 1845. Catalogue of Lizards in the British 
Museum, London. 

Greer, A. E. 1967. A new Generic Arrangement for 
Some Australian Scincid Lizards. Breviora 267 

Loveridge, A., 1934. Australian Lizards in the Mu- 
seum of Comparative Zoology, Cambridge, Mas- 
sachusetts. Bull. Mus. comp. Zool. Harw 11' 6 

Mittleman, M. B., 1952. A Generic Synopsis of the 
Lizards of the Subfamily Lygosominae. Smith- 
son, misc. Colin. 117: 17. 

Storr, G. M. In press. The Genus Hemiergis Lacer- 
tilia, Scincidae in Western Australia. 

Werner, F., 1910. Fauna Siidwest-Aust. 2: 480 



THE ENDEMIC AUSTRALIAN LIZARD GENUS MORETHIA 
(SCINCIDAE; LYGOSOMINAE) IN SOUTHERN AUSTRALIA 

By P. A. Rawlinson 

Zoology Department, La Trobe University, Bundoora, Victoria, 3083 

Abstract 

The taxonomy and status of the five southern Australian species of Morethia are discussed 
and lectotypes are nominated to stabilize the nomenclature. Details of the species distributions 
are provided and the ecology, reproduction, generic relationships and phylogeny of the species 
are briefly mentioned. 



Introduction 

Boulcnger (1887) carried out the first 
major revision of the family Scincidae in the 
third volume of his Catalogue of Lizards in 
the British Museum (Natural History). In the 
Preface to this volume Dr A. Gunther stated: 
'I feel confident that it will give a fresh 
impluse to the systematic study of lizards, 
and serve as the standard work for many 
years to come 1 . Gunther's confidence was gen- 
erally well placed and Boulenger's Catalogues 
became the standard reference works. How- 
ever, in some groups rather than give a fresh 
impulse to systematic studies, Boulenger's 
concepts caused a stagnation that lasted more 
than half a century. One such group was his 
large and clumsy genus Lygosoma with it's 
eleven subgenera, and another was the genus 
Ablepharus into which Boulenger placed all 
skinks with an immoveable transparent lower 
eyelid (the 'ablepharine' eye). These genera 
were long recognized to be polyphyletic, but 
it wasn't until Mittleman (1952) erected the 
subfamily Lygosominae and revised the in- 
cluded genera that a new stimulus was pro- 
vided. Mittleman included the species Boul- 
enger had placed in Ablepharus in the Lygos- 
ominae and broke up the genus. Since that 
time work on the higher taxa of Lygosomine 
skinks has made rapid progress and the 
species Boulenger included in the genus Ab- 
lepharus have been reclassified. 

The Australian skinks which were included 
in Boulenger's definition of the genus Able- 
pharus have been dealt with in two main 
papers. Greer (1967) convincingly demon- 
strated the artificial nature of the 'ablepharine' 
eye taxonomically and phylogenetically when 



he united a group of closely related 'able- 
pharine' and 'non-ablepharine' skink species 
in the genus Lerista. Subsequently Fuhn 
(1969) separated the Australian 'ablepharine' 
skinks into nine groups, one of which was 
the genus Morethia Gray, 1845. Fuhn sep- 
arated the genus Morethia on the basis of 
skull morphology and in doing so successfully 
placed a natural group of closely related 
species into one genus. Morethia is now re- 
cognized to be an endemic Australian genus 
which is not clearly related to any non -Aus- 
tralian genus (Storr 1972) and even its 
Australian relationships are unclear (Rawlin- 
son 1974). The genus presently consists of 
six described species, two described subspec- 
ies and an undescribed 'race' (Storr 1972). 

The Morethia species and subspecies (and 
therefore the genus) can be divided into two 
geographical and evolutionary groups: a 
'northern' group centered in the arid and 
semi-arid tropical areas of Australia; and a 
'southern' group centered in the arid and 
semi-arid temperate areas of Australia. The 
'northern' group consists of: M. taeniopleura 
taeniopleura (Peters 1874) found in N and 
E Queensland, which reaches to 27 i° S but 
is centered N of the Tropic of Capricorn 
23i° S; M. taeniopleura ruficauda (Lucas 
and Frost 1895) found in N Northern Territ- 
ory and N Western Australia, which reaches 
to 25i° S but is centered N of the Tropic of 
Capricorn ; M. taeniopleura exquisita Storr 
1972 found in NW Western Australia, which 
reaches to 25° S but is centered N of the Tropic 
of Capricorn; and an undescribed race of 
M. taeniopleura recorded from the N of the 
Northern Territory by Storr (1972). These 



27 



28 



P. A. RAWLINSON 



'northern' taxa are allopatric and appear to 
be mutually exclusive (Storr 1972). 

The 'southern' group consists of the re- 
maining five described species and it appears 
to have radiated more widely than the 'north- 
ern' group. Three of the five 'southern' spec- 
ies, M. adelaidensis, M. butleri and M. ob- 
scura, although reasonably widely distributed 
are restricted to arid and semi-arid areas S 
of 271° S; the fourth species, M. lineoocel- 
lata, is virtually restricted to the SW coast 
of Western Australia but it extends above the 
Tropic of Capricorn to about 20i° S; the 
fifth species, M, boulengeri, is widely distii- 
buted across S Australia and it extends just 
above the Tropic of Capricorn to about 22i° 
S in S Queensland. Of the 'southern' taxa, 
M. boulengeri and M. butleri are allopatric 
and appear to be mutually exclusive as do 
M. lineoocellaia and M. obscura; M. butleri 
and M. adelaidensis are largely allopatric but 
overlap in SE Western Australia; and M. 
adelaidensis, M. boulengeri and M. obscura 
overlap widely across S Australia. Thus it 
can be seen that the Tropic of Capricorn 
forms a boundary between the 'northern' and 
'southern' groups of Morethia and it is the 
five 'southern' species as defined above which 
are dealt with in detail in this paper. 

Two recent local revisions of the genus 
Morethia included the five 'southern' species. 
The first revision, published by Smyth (August 
31, 1972) was of the South Australian species 
but it also included all specimens of Morethia 
in the South Australian Museum. Storr pub- 
lished the second revision (November 3, 1972), 
which was of the Western Australian species 
but it also included all specimens of Morethia 
in the Western Australian Museum. Unfor- 
tunately Symth's and Storr's papers contain 
some conflicting interpretations and neither 
author examined the types of Morethia lineo- 
ocellaia (Dumeril and Bibron 1839) which 
was the first of the Morethia species described 
and hence the most important taxonomically. 
The present paper deals with the 'southern' 
Morethia species in Queensland, New South 
Wales and Victoria, and as the author has 
examined all relevant type specimens, the 



opportunity to correct the conflicts between 
Smyth's and Storr's papers is taken. Also, as 
the author has examined and identified all 
Morethia specimens in the Queensland Mus- 
eum (QM) Brisbane, the Australian Museum 
(AM) Sydney, and the National Museum of 
Victoria (NMV) Melbourne, detailed lists of 
these specimens are provided under the ap- 
propriate headings below to complement the 
data in Smyth and Storr. As there is no 
Morethia material in the Tasmanian Museum, 
Hobart, or the Queen Victoria Museum, Laun- 
ceston, the data in Smyth's and Storr's papers 
and the present paper represents a complete 
listing of the 'southern' Morethia specimens 
held in the Australian state museums. 

Genus Morethia Gray, 1845 

Morethia Gray, J. E., 1845, Catalogue of lizards: 65. 

TYPE SPECIES; Morethia anomalus Gray, 1845, 
Ibid.: 65 — Ahlepharus lineoocellatus Dumeril, 
and Bibron, 1839, Erpetologie Generate 5: 817. 

Remarks; Smyth (1972) incorrectly listed 
Ablepharus lineoocellatus Dumeril and Bibron 
1839 as the type species of the genus. Storr 
(1972) listed Morethia anomalus Gray 1845 
as the type species by monotypy and in the 
same paper designated M . anomalus as a jun- 
ior subjective synonym of A. lineoocellatus. 
Diagnosis: Small skinks (snout-vent length 17- 
56 mm); an 'ablepharine' eye i.e. lower eyelid 
an immoveable transparent disc fused to the 
eye surface; Frontoparietals and interparietal 
fused into a single large shield; parietals con- 
tact along midline; supranasal and postnasal 
scales present but may be fused to each other 
or to nasal scale; frontonasal in broad contact 
with the rostral; frontal much larger than the 
prefrontals; prefrontals rarely in contact; four 
supraoculars, second the largest, first and sec- 
ond contact the frontal, second third and fourth 
contact the frontoparietal-interparietal shield; 
one pair of nuchal scales; seven (occasionally 
eight) upper labial scales, the fifth largest 
and completely subocular; eight to ten preanal 
scales, central four slightly enlarged; limbs 
pentadactyl; digits not elongate, 14-27 lamellae 
under the fourth toe; body scales smooth, 
moderately large, 24-34 rows at midbody; 
external ear opening obvious. 



THE ENDEMIC AUSTRALIAN LIZARD 29 

In the descriptions of Morethia species and — Storr, 1972, /. R. Soc. W. Aust. 55: 73-79. 

specimens below, scalation details consistent Flgs " ls 2 ' 

with the generic description above are not Lectotype: Smyth (1972): ZMB 4733, Zoo- 
repeated, logisches Museum der Humboldt Universitat 

zu Berlin. Locality: Adelaide. Collector: 

KEY TO THE SOUTHERN AUSTRALIAN Schomhun* No other data 

SPECIES OF MORETHIA 2>cnomDurgk. JNo otntr data. 

1 . Subdigital lamellae acutely keeled, uni- D scnp , lor L u e< f + m / , /' u c ,, . 

. °. , . . . a . Remarks: The lectotype, selected by Smyth in 

cannate to tricannate; five or srx sup- ln71 +u , '/ i it / i .. 

.,- . ' * 9 1971, was the largest of three syntypes under 

c i. a- u i i 11 xi ■ " ■•■•'' ' this catalogue number and the paralectotypes 

Subdigital lamellae smooth or obtuselv , & u r u 4*kkt* 

, . r .... „ have now been given new numbers, ZMB 

keeled; six supracihanes 3. .^^ n ~ , , to - , . , 

n t-,. ' ... \ .. A , . , r . 42872-73, data as for lectotype. 

2. Five supracilianes, the third, fourth c +u nn7 ^ , c , A^nn\ u fu 

* ,J t ' . i j Smyth (1972) and Storr (1972) both coni- 

and fifth largest, subequal, and pen- . J A n / , /107 ^ c + u « ~ 

t t j i t, J th l mented on Peters (1874) use of the name 

i j- -x i 1 ii ■ • ' Ablepharus (Morethia) anomalus adelaidensis 

subdigital lamellae unicarmate or a -<.u a a n +~ > a • *~~ a~ 

. . " . . . , , . , and neither considered Peters description ade- 

tncarinate M. adelaidensis . TT c ., . , , „ . , 

. .,, . .- ~ ^ ., - quate. However, Smyth considered Peters use 

Six .supracilianes, the first the largest Qf ^ mm& constituted a valid indication of 

and the remainder forming a deereas- a d ^^ ^ namfi tQ hi aM ^ 

ing series, junctions of supracilianes cQrdi sekcted Qne of pet£rs , three 

with supraocular linear or slightly frQm ^ BerM MmQum as k s 

curved, supracilianes do not pene rate ho did no[ ^.^ pete use Qf flie 

between supraoculars; subdigital lam- name constituted a vaIid indication of a specie s 

ellae unicarmate .... . . . M. feffen ^ he refused tQ ^^ ^ name tQ p eters> 

3. First and third supracihanes largest, bu{ he did no{ formall ^ k ^ & mmen 
fourth much smaller than third, fourth, mdum Boul (1887) had ascribed the 
fifth and sixth successively smaller name ^MaWe,^ to Peters when redescribing 

r . .,. , M - boulen 8 en the 'variety' as Ablepharus lineoocellatus C var. 

First supracihary never largest, fourth adelaidensis, and there is every chance he ex- 

not smaller than third ■;•■.■■■-* amined Peters' types for he stated in the In- 

4. Third, fourth and fifth supracihanes traduction to Volume 3 of his Catalogue 'With 
largest, subequal and penetrate deeply ^ object of renderine th e account of the 
between the supraoculars, sixth much Lacertidae and Scincidae more perfect, I have 
smaller than fifth; supranasa often deyoted a mmth tQ the examination of the 
fused to nasal . M. Imeoocellala imens in the Berlin Museum'. Storr re- 
Fourth supracihary largest, fourth, fifth ded Boule er as the pro er authority for 
and sixth form a rapidly decreasing ^ name adel ^ demis and according iy selected 
series, third and fourth penetrate deep- a lectot from the spec i m ens described by 
ly between supraoculars; supranasal B oulenger in the collection of the British 
always separate from nasal. . . M. obscura Museum of Natural History m London: BM 

Morethia adelaidensis (Peters 1874) NH 64.10.27.9; Locality: South Australia; 

.p. .J -. Collector: G. Krefft; no other data. Smyth had 

already examined the specimen which Storr 

Ablepharus (Morethia) anomalus adelaidensis Peters, „ n u sprllipnt i u Hp<;ianaterl as lerfntvrvp anH iHpn- 

1874, Mher. Preuss. Akad. Wiss.: 375-376. subsequently designated as lectotype and lden- 

Ablepharus lineoocellatus C var. adelaidensis Boulen- tified it as M. adelaidensis. The present author 

ger, 1887, Catalogue of the Lizards in the British has re _ ex amined Storr's lectotype and confirms 

Museum (Natural History). 3: 349. . . , *' t , , 

Ablepharus lineoocellatus (part) Zietz, 1920, Rec. S. it is conspecific with the lectotype selected by 

Aust. Mus. 1: 220-221. Smyth (ZMB 4733). 

M Mutl6 ad iftits. hT' 1972 ' ^ S ' Smyth's designation of the lectotype is cor- 



30 



P. A. RAWLINSON 



rect and must be upheld. Storr's action would 
have been valid if Peters' use of the name 
adelaidensis constituted a nomen nudum, but 
Storr did not state this was the case and articles 
11., 12. and 16. of the International Code 
for Zoological Nomenclature ( 1 964 ) show 
Peters' use of the name constituted a valid 
indication of the species. It should be noted 
that Boulenger himself ascribed the name 
adelaidensis to Peters, so both authors were 
referring the name to specimens Peters had 
examined in the Berlin Museum and there 
is a great probability that Boulenger actually 
examined Peters' types. Smyth located these 
specimens and designated one as lectotype, and 
as Smyth's paper was published on August 31, 
1972 and Storr's paper was published on Nov- 
ember 3, 1972, Smyth's designation predates 
Storr's. 

Diagnosis: Five supraciliaries, the third, fourth 
and fifth the largest, subequal, and all penet- 
rate deeply between the supraoculars. Sub- 
digital lamellae acutely keeled, unicarinate or 
tricarinate. Palmar tubercles elongate and 
apically rounded. 

Description: Snout-vent length 17-60 mm, 
mean 46 1 mm. Total length adults with intact 
tails 104-138 mm, mean 118-8 mm. Intact 
tail 120-172% of snout-vent length. Suprana- 
sals present, widely separated. Postnasals pre- 
sent but often fused to supranasals. Prefrontals 
narrowly separated. Frontonasal wider than 
long. Frontal longer than wide. One to three 
ear lobules, usually hidden by projecting prc- 
auriculars. Midbody scales in 26-34 rows (us- 
ually 28 or 30), mean 29-1. Lamellae under 
fourth toe 16-24, mean 19-7. 
Colour: Olive-grey to olive-brown dorsal sur- 
face, often tinged with red-brown. Small black 
spots on back which tend to form broken 
lines. Pale dorsolateral stripe occasionally pre- 
sent on trunk. Broad dark brown to black 
upper lateral stripe strongly speckled with 
lighter markings runs from head onto tail. 
Wavy edged interrupted white mid-lateral line 
runs from upper labials, through ear, above 
forelimb and along trunk to hindlimb, usually 
margined below by a speckled brown band. 
Ventral surface unmarked, white. Males in 



breeding condition develop an orange colour 
all around the edges of the ventral surfaces 
which extends onto the inside surfaces of both 
fore and hind limbs and is particularly prom- 
inant around the vent and anterior part of the 
tail. 

Distribution: Arid and semi-arid-areas of SW 
Queensland; SW New South Wales; NW Vic- 
toria; NE to S South Australia; and SE Western 
Australia. (Figure 1.). 




Fig. 1. M. adelaidensis 



Literature Records: See lists in Smyth (1972) 
and Storr (1972). 

Specimens Examined: Western A usiralia: 
(NMV) Western Australia, R 963; D 1013; 
D 1390: 15 mis E of Caiguna, D 44857: South 
Australia: (AM) South Australia, 4739: 
Fisher, Nullabor Plain, R 7274: (NMV) Cen- 
tral Australia, D 1181; D 1183-4: Gawler 
Ranges, D 2464-5: Ooldea Well, Overland 
Railway, N of Fowlers Bay, D 2473: Overland 
Railway, between Ooldea Well and Fowlers 
Bay, D2758: Overland Railway to Western 
Australia, D 3059; D3109; Lake Eyre, D 
3103; D 3128; D 3135: Lake Harry, Birdsville 
Track, D 15027: Lake Wangary, Eyre Penin- 
sula, D 15054-7; D 15059-60: 3 km NW of 
Poonindie, D 15078: Tumby Bay, Eyre Pen- 
insula D 15078-80; D 15217: 18i km S of 
Maitland, Yorke Peninsula, D 15093-7: Cop- 
ley, D 15847-8: Benagerie Station, D 41509, 
D 41525: Innamincka area, D 41606-7: 
New South Wales: (AM) Moloch, R6445 



THE ENDEMIC AUSTRALIAN LIZARD 



31 



(3 specimens): Victoria: (NMV) Grampians, 
D 1090': 16 km W of Nowingi, D 14652: 13 
km S Stawell, D 15064: Kerang, D 15163: 32 
km S of Kaniva, D 15174-5: 29 km N of Swan 
Hill,D 15177: 10 km W of Nowingi, D 18106. 

Morethia boulengeri (Ogilby 1890) 

(Hg. 2) 

Ablepharus boulengeri Ogilby, 1890, Rec. Aust. Mus. 

1: 10-11. 

Zietz, 1920, Rec. S. Aust. Mus. 1: 220. 

Ablepharus lineoocellatus anomalus (part), Loveridge, 

1934, Bull. Mus. Comp. ZooL 77: 377-378. 
Morethia boulengeri Smyth, 1972. Rec. S. Aust. Mus. 

16: 1-14. Figs. 2, 6. 
Storr, 1972, /. R. Soc. W. Aust. 55: 73-79. 

Figs. 1, 2. 

Lectotype: Smyth (1972): AM R 690, Aus- 
tralian Museum, Sydney. Locality: Brawlin, 
New South Wales (34° 44' S 148° 02' E). 
Collector: H. J. McCooey. No other data. 
Description: See Smyth (1972). 
Remarks: Smyth (1972) noted that Ogilby 
incorrectly recorded a separate interparietal for 
the type. Storr (1972) listed Cootamundra, 
New South Wales (10 km N of Brawlin) as 
the type locality, which is the locality given 
for the specimen in the Australian Museum 
Register. 

Diagnosis: Six supraeiliaries, the first and third 
the largest; the fourth much smaller than the 
third; and the third, fourth, fifth and sixth 
are successively smaller. Subdigital lamellae 
smooth or obtusely unicarinate. Palmar tuber- 
cles rounded. 

Description: Snout-vent length 27-55 mm, 
mean 44-6 mm. Total length adults with intact 
tails 102-121 mm, mean 109 mm. Intact tail 
125-177% of snout-vent length. Supranasals 
present, widely separated. Postnasals present 
but often fused to supranasals. Prefrontals 
separated. Frontonasal wider than long. Front- 
al longer than wide. Seven upper labials norm- 
ally, rarely eight but the fifth always largest 
and entirely subocular. Two to four obtuse 
ear lobules (usually two). Midbody scales in 
25-32 rows, mean 29-8. Lamellae under the 
fourth toe 15-23, mean 19-6. 
Colour: Olive-grey to brown dorsal surface, 
dorsal scales have 2-5 (usually 3) fine diver- 



ging black lines which are often expanded and 
merge into black spots or streaks that are 
usually distributed irregularly, but may be 
organized into interrupted lines or streaks. A 
well-defined broad black upper lateral stripe 
runs from the eye back to the hindlimb where 
it becomes broken up. A prominent moder- 
ately wide pure white mid-lateral stripe begins 
on the upper labials and runs under the eye 
above the forelimb and along the trunk to the 
hindlimb. Usually there is also a narrow irreg- 
ular black lower lateral stripe below the mid- 
lateral stripe. Ventral surface unmarked silver 
white. Adult males in breeding condition dev- 
elop a bright orange throat. The tail of juv- 
eniles is red-orange, not pale fawn as recorded 
by Smyth (1972), adults have brown tails. 
Distribution: Drier areas of S Northern Terri- 
tory; S Queensland; New South Wales; NW 
and N Victoria; South Australia; and SE 
Western Australia. (Figure 2.). 




Fig. 2. M. boulengeri 



Literature Records: See lists in Smyth (1972) 
and Storr (1972). 

Specimens Examined: Northern Territory: 
(NMV) Illamurta, James Range, D 473: 
Queensland: (AM) Retro Station, Capella, R 
12099 (2 specimens); R 12100 (2 specimens); 
R 12114 (3 specimens): Mungindi, R 15073; 
R 19083: Cunnamulla, R 17121; R 18464: 
Gilruth Plains, Cunnamulla, R 20662-3: 8 km 



32 



P. A, RAWLINSON 



S of Claremont, R 21312: Glenmorgan, R 
21317: (NMV) Queensland, D 104: 5 km 
W of Amiens, D 9384: (QM) Bell, Darling 
Downs, J 2305-9: Retro Station, Capella, 
J 6232; J 15768: 24 km W of St. George, J 
10456-7: 69 km S of Blackall, J 11564: 
Wivenhoe Bridge, Brisbane River Valley, 
J 1 1853: 24 km N of Mitchell, J 11854: Bry- 
den, J 1 1855: Green swamp, 45 km E of Roma, 
J 11856; J 11954: Murphy's Lake, Taroom, 
J 11858: 6 km S of Ferndalc, J 11859-60: 
Wivenhoe Bridge, 8 km N of Wivenhoe, 
J 11862-3: 10 km W of Westgrove HS., 
NW of Gympie, J 11864: li km W of West- 
grove HS., NW of Gympie, J 11865: Win 
dorah, J 11866-8: Waratah Station via Cun- 
namulla, J 11869-71: Alum Rock Station via 
Amiens, J 11872: 16 km SE of Capelta, 
J 11953: 8 km N of Stanthorpe, J 11955: 
Goondiwindi, J 13682: Jandowae, J 13773; 
J 13775-6; J 13778-81: Gilruth plains Field 
Station, Cunnamulla, J 14353: Bathcaston Sta- 
tion, 128 km NW of Marlborough, J 15635-7: 
35 km N of Augathella, J 21609: 88 km 
NW of Tamboo, J 21610; Binga Station via 
Cunnamulla, J 21946: Amiens near Stan- 
thorpe, J 22316: Jollys Falls via Stanthorpe, 
J 22476: South Australia (AM) Port Lincoln, 
R 4764: (NMV) Purnong, D 1216: Overland 
Railway to Western Australia, D3058: Why- 
alla, D 7743; D 7979-80: Alligator Gorge, Flin- 
ders Range, D 15220-2: Wertalona, D 41542: 
New South Wales (AM) Cootamundra, 
R 687-91; R 950: Brawlin, R 823; R 825-6: 
Narrabri, R 1042; R 1058: Western New South 
Wales, R 1073-4: Nidgera Ck., near Coot- 
amundra, R 1745: Dubbo, R 1750-61: Nar- 
romine, R 1780: Boggabri, R 2023; R 2025; 
R 4169: near Liverpool, R 2174-5: Gunta- 
wang, R 3976: Nevertine to Warren, R 4871: 
Buddah Lakes, Trangii, R 5447: Penrose 
Farm, Bathurst, R 6268: Moloch, R 6443: 
Darling R., between Bourke and Wilcannia, 
R 6457: Belubula, R 8077: Jaffa, Ashford 
Downs, R 9413: Yaneo district, R 10646: 
Yaneo Agricultural School, R 10648: Walcha, 
R 10769: Cuddie Springs, Brewarrina, R 
11006: Bullerana, Moree, R 11054: Pilliga, R 
1 1591 : Bringagee, R 13432: Mootwingie 



Watcrholes, R 14678-9: Warrumbungle Moun- 
tains, R 14973; R 18921; R 21076: Quam- 
bone, R 15120; R 18722; R 18728; R 18753- 
4; R 18783: 21 km W of Byrock, R 15212: 
24 km W of Byrock, R 15621: Moulamein, 
R 15841: Brewarrina, R 17674; R 18727: 
Lake Narran, R 17658; R 17711: Bourke, 
R 17886: Tocumwal, R 18143-5: Balranald, 
R 18146-8: Murray R., Moama, R 20357: 
Nymangec, R 20572; R 20642-3: 72 km W 
of Cobar, R 27409-10: Round Hill Fauna 
Reserve, between Lake Cargelligo and Mt. 
Hope, R 27825; R 27834; R 27839-40; R 
27862-4; R 27904; R 29669: Old Harbour 
Lagoon, near Eumungie, R 28021-2; R 280 
24-6: between Tilpa and Barnato, R 28093: 
Coonabarabran, R 28095: Duck Ck., 48 km 
W of Warren, R 28160: 67 km S of Gilgunnia, 
Eubalong Rd., R 28341; R 28347-8: 32 km 
S of Cobar, R 29560: Yalgorgrin State Forest, 
11 km ex. Gilgandra, R 29645: Lachlan R., 11 
km N of Lake Cargelligo, R 30385-7: 64 km 
S of Gilgunnia, Mt. Hope to Eubalong Rd., 
R 30391 : Moonabi Lookout, Moonabi Ranges, 
R 31741-2: 37 km W of Armidale on Bun- 
darra Rd., R 31769; R 31785: 24 km W of 
Gilgandra, R 33087-98; R 33100-6: (NMV) 
Finley, D 152: Brawlin, D 749-51: Nyngan, 
D 10591: Gin Gin, D 10603: Tor Downs, SW 
of Menindee, D 10610-1: Moorna, D 12167: 
16 km N of Albury, D 14641-2: Savernake 
Station, Savernake, D 15043-8; D 15061-3: 
Boat Rock Hill, Savernake, D 15065-6: 
8 km S of Corowa, D 15067-77: 6i km SW 
of Savernake, D 15081: Mulwala, D 15086-7: 
Warrumbungle National Park, D 15091: 6i 
km S of Morandah, D 15092: 14i km NE of 
Urana, D 15130: 3 km NE of Jerilderie, D 
15131: 14i km NE of Jerilderie, D 15132: 
3 km N of Tocumwal, D 15133: Tocumwal, 
D 15134: 10 km N of Cowra, D 15135: 
Boree Creek, D 15136-8; D 15142-5: 21 km 
SE of Jerilderie, D 15139-41: Sloane, D 15146: 
13 km SE of Jerilderie, D 15147: 10 mis 
N of Barham, D 15178-92: 90 km W of 
Wentworth, D 15850: 24 km NNE of Went- 
worth, D 40180-1: Victoria (AM) Browns 
Plains, R 3988: (NMV) Murray River, D 
917: Ouyen, D 810-1; D 950: Mildura, D 



THE ENDEMIC AUSTRALIAN LIZARD 



33 



848; D 15218-9: Victoria, D 945: Elmore, 
D 1002: Bright, D 1065; D 1067; D 1069: 
Tcmplestowe (in error) D 1237: Dimboola, 
D 1724; D 2207: Goulburn Valley, D 1808: 
Glenrowan, D 1819; D 2517: near Melbourne 
(in error) D 2508: Cowangie, D 2717: Gram- 
pians, D 3315-6: Kewell, D 3321-3: Red 
Cliffs, D 7966; D 7968: Sea Lake, D 9104-5: 
Pink Lakes, Linga, D 10713: Birthday Tank, 
Sunset Country, D 11185-6; D 33305; D 
44865; D 47407: Wood Wood, D 13948-9: 
Taminick Gap, Warby Ranges, D 14553-63; 
D 14573-5: Warby Ranges, D 14610: Speci- 
men Hill, Byawartha, D 14635: 8 km S of 
Kiata, Little Desert, D 14911-5; D 14984: 8 
km SW of Kiata Little Desert, D 14953: 8 km 
SW of Natimuk, D 15042: Walpeup, D 150 
49-50: 13 km N of Underbool, D 15051-2: 
10 km SE of Wycheproof, D 15053: 13 km 
E of Yarrawonga, D 15082: 6i km S of 
Nathalia, D 15083: 13 km S of Yarrawonga, 
D 15084-5: Kenmere, D 15088: 16 km SW 
of Nathalia, D 15089-90: Tutye, D 15097-103: 
Pine Plains, D 15104-5: 13 km E of Patche- 
wollock, D 15106: Axedale, D 15108: 4 km 
E of Mildura, D 15109: Wemen, D 15110: 

4 km NE of Wemen, D 15111: 4 km 
NE of Mildura, D 15112: 7 km WSW of 
Leitchville, D 15113-5: 35 km NW of Nyah, 
D 15116: Hattah, D 15117: Mildura Airport, 
D 15118-29: Lindsay Point Station, D 15148: 

5 km E of Neds Corner Station, D 15149-53; 
D 15157-8: 6i km E of Neds Corner Station, 
D 15154-5: 6£ km NW of Neds Corner 
Station, D 15156: Neds Corner Station, D 
15159: Kerang, D 15160-2; D 15164: 5 km 
N of Eaglehawk, D 15165-6: 6i km NE of 
Wahgunyah, D 15167-72: 6i km E of Tal- 
garno, D 15173: 29 km N of Swan Hill, D 
15176: Nyah, D 15193: 3 km E of Hattah, 
D 15194-206: Tower Hill, Grampian Ranges, 
D 15208: 5 km NW of The Crater, Little 
Desert, D 15210-3: 6i km N of The Crater, 
Little Desert, D 15209: Manangatang, D 
15214-6: Tiger Hill via Tatong, D 17710-7: 
between Perry and Birthday Tanks, Sunset 
Country, D 18064-6: H km E of Birthday 
Tank, Sunset Country, D 18076: 7 km W of 
Nowingi, D 18102-4: Tank, 6i km E of 



Birthday Tank, Sunset Country, D 18165: 
SW bank of Rocket Lake, Sunset Country, 
D 1 8 1 68 : between Monkeytail and Perry 
Tanks, Sunset Country, D 18173: Stockyards 
between Perry and Birthday Tanks, Sunset 
Country, D 18175-9: Stockyards, Sunset Tank, 
Sunset Country, D 18184-5: 5 km NE of 
Sunset Tank, Sunset Country, D 18233: 1 km 
E of Sunset Tank, Sunset Country, D 18234: 
24 km E of Birthday Tank, Sunset Country, 
D 33304: 1 km S of Millewa South Bore, 
Sunset Country, D 33347: 14i km NE of 
Millewa South Bore, Sunset Country, D 
33350-1: Millewa South Bore, Sunset Country, 
D 33354: Pound Bend, Wemen via Robinvale 
D 33358-60: Kulkyne Freehold, D 33528-30: 
6i km SE of Pine Plains, D 33536: 5 km E 
of Broughton, Nhill Rd., D 33638-43: H km 
N of Freuds Plain, Wyperfeld National Park, 
D 38799: Main Gate, Wyperfeld National 
Park, D 38800-5: 3 km W of Yanac, D 38818: 
Sunset Tank, Sunset Country, D 38853; 
D 38860-1: 3 km E of Perry Tank, Sunset 
Country, D 38856: 3 km N of Bendigo, 
D 44834-6: Mildura, D 40141-2: 11 km 
WNW of Annuello, D 47343-4: 11 km S of 
Boundary Bend, D 47354-6: 

Morethia butleri (Storr 1963) 
(Fig. 3) 

Ablepharus butleri Storr, 1963, W. Aust. Nat. 9: 

46-47. 
Morethia butleri Smyth, 1972, Rec. S. Aust. Mus. 16: 

1-14. Fig. 3. 
Storr, 1972, J. R. Soc. W. Aust. 55: 73-79. 

Figs. 1, 2. 

Holotype: WAM R 20615, Western Australian 
Museum, Perth. Locality: Leonora, Western 
Australia, 28° 52' S., 121° 23' E. Collectors: 
G. M. Storr and R. E. Moreau. 
Description: See Storr (1963). 
Diagnosis: Six (rarely seven) supraciliaries, 
the first the largest and the remainder forming 
a decreasing series; junction of supraciliaries, 
with supraoculars linear or slightly curved. 
Subdigital lamellae acutely keeled and unicari- 
nate. Palmar tubercles apically acute. 
Description: (After Smyth (1972) and Storr 
(1972)) Snout-vent length 25-56 mm. Intact 
tail 134-169% of snout-vent length. Suprana- 



34 



P. A. RAWLINSON 



sals present, widely separated. Postnasals pre- 
sent but often fused to supranasals. Prefrontals 
separated. Frontonasal wider than long. Front- 
al longer than wide. Two to five ear lobules 
(usually two or three). Midbody scales in 26- 
31 rows (usually 28 or 30), mean 28-9. 
Lamellae under fourth toe 19-27, mean 22-4. 
Colour: Dorsal surface dark olive-green to 
olive-brown sometimes flecked with black, but 
usually unmarked. Broad black upper-lateral 
stripe may be distinct and well developed or 
virtually absent except anteriorally. The white 
mid-lateral stripe is equally variable, it may 
be distinct and well developed or only readily 
distinguishable anteriorally. Lips dark spotted. 
Storr (1972) records that the tail (as with M. 
boulengeri) is red in juveniles and brown in 
adults. 

Distribution: Arid and semi-arid parts of S 
Western Australia and possibly SW South 
Australia S of 27° 30' S. Not known from 
any other part of Australia. (Figure 3.). 




Literature Records: See lists in Smyth (1972) 
and Storr (1972). 

Specimens Examined: South Australia (NMV) 
Overland Railway to Western Australia, D 
2654: No Data (AM) R 38652. 



Morcthia lineoocellata (Dumeril and 
Bibron 1839) 

(Fig. 4) 

Ablepharus lineoocellatus (part) Dumeril and Bibron, 

1839 Erpetologie Generate 5: 817. Paris. 
Morethia anomalus (part) Gray, 1845, Catalogue of 

lizards: 65. 
Ablepharus lineoocellatus A var. lineoocellatus Bou- 

lenger, 1887, Catalogue of the Lizards in the 

British Museum (Natural History) 3: 348-349. 
Ablepharus lineoocellatus B var. anomalus (part) 

Boulenger, 1887 Ibid. 3: 48-349. 
Ablepharus lineoocellatus Zietz, 1920, Rec. S. Aust, 

Mus. 1: 220-221. 
A blepharus lineoocellatus lineoocellatus Loveridge, 

1934, Bull. Mus. Comp. Zool. 11: 111. 
Morethia lineoocellata Fuhn, 1969, Z. Zool. Svst. 

Evolutionsforsch. 1: 67-76. Fig. 7. 
Morethia lineoocellata Storr, 1972, J. R. Soc. W. Aust. 

55: 73-79. Figs. 1, 3. 

Type Series of Ablepharus lineoocellatus, 

Dumeril and Bibron, 1839 
Remarks: Five specimens (not four as listed 
by Guibe, 1954) in the Museum National 
d'Historie Naturelle, Paris: MNHP 3092 (old 
number 3101). Locality: Nouvelle Hollande. 
No other data. Two distinct species are repre- 
sented in the type series which was not exam- 
ined by either Smyth or Storr. Three of the 
syntypes are referable to M. lineoocellata 
sensu Storr. The first of these syntypes (the 
largest) fits Stores definition of M. lineoocel- 
lata in all respects, this specimen is designated 
as lectotype below. The second syntype has 
a partially separated supranasal scale which 
is characteristic of Stores M. obscura, but not 
exclusive of M. lineoocellata, this specimen 
otherwise fits Storr's definition of the latter 
species. The third syntype is aberrant, it has 
five supraciliaries owing to the fusion of the 
first and second, and it fits Storr's definition 
of M. obscura in one respect, for the third 
last supraciliary is the largest and the last three 
supraciliaries form a rapidly decreasing series. 
However, this specimen lacks supranasal scales 
which are characteristic of M. obscura. The 
remaining two syntypes are conspecific and 
distinct from either M. lineoocellata or M. 
obscura; both specimens are readily referable 
to Menetia greyi Gray 1845 (see below). 

In order to restrict the use of the name 
Morethia lineoocellata and preserve Storr's 



THE ENDEMIC AUSTRALIAN LIZARD 



35 



nomenclature, the first syntype listed above 
which fits Store's definition exactly is selected 
as lectotype here. 

Lectotype: MNHP 3092 (Old No. 3101), 
Museum d'Historie Naturelle, Paris. 
Locality: Nouvelle Hollande (^Australia) . 
No other data. 

Description: Snout-vent length 42 mm. Tail 
(intact) 51 mm, 121% of snout-vent length. 
Length of forelimb, 12 mm; length of hindlimb, 
18 mm. Snout-axilla, 10 mm; axilla-groin, 
24 mm. Six supraciliaries, third, fourth and 
fifth subequal in size and penetrating deeply 
between the supraoculars; sixth supraciUary 
the smallest. Supranasals fused to nasals. Post- 
nasals present. One ear lobule. Seven upper 
labials, the fifth largest and entirely subocular; 
six lower labials. One pair of nuchals. Eight 
preanals, four slightly enlarged. Smooth scales, 
26 rows round midbody. Subdigital lamellae 
smooth, 19 under the fourth toe. Palmar tuber- 
cles apically rounded. 

Colour: Dorsal surface olive-brown with two 
rows of rather indistinct black and white 
ocelli. Dorsolateral row of indistinct black and 
white ocelli. Black upper lateral band runs 
from nostril through the eye, above the fore- 
limb and along trunk to tail. Distinct white 
mid-lateral stripe runs from upper lip through 
ear and above fore and hind limbs to tail. 
Black lower lateral band runs below white 
mid-lateral stripe, merges into grey lower 
lateral region. Ventral surface pure white un- 
marked. 

Paralectotypes: Four specimens also under 
MNHP 3092 (Old No. 3101), Museum d'His- 
torie Naturelle, Paris. 
Locality: Nouvelle Hollande. No other data. 
Paralectotype (a): Snout-vent length 35 mm; 
tail (regrown) 41 mm; length of forelimb 11 
mm; length of hindlimb 16 mm; snout-axilla 
13 mm; axila-groin 20 mm. Scalation as for 
lectotype except: supranasal only partially 
separated from nasal; 24 rows of scales round 
midbody; 18 lamellae under the fourth toe; 
ten preanal scales, four slightly enlarged. 
Colour as for lectotype except: each of the 
two dorsal rows of black and white ocelli have 
fused to give continuous white lines margined 



by broken black lines on the trunk ad tail; 
and on each side the dorsolateral row of 
ocellations have also fused to given a con- 
tinuous white dorsolateral stripe margined 
above and below on the trunk by black. This 
specimen is conspecific with the lectotype. 
Paralectotype (b): Snout-vent length 41 mm; 
tail (regrown) 19 mm; length of forelimb 11 
mm; length of hindlimb 17 mm; snout-axilla 
9 mm; axilla-groin 20 mm. Scalation as for 
lectotype except: five supraciliaries owing to 
the fusion of the first and second, second and 
third the largest, only the second and third 
penetrate deeply between the supraoculars, 
and the third, fourth and fifth form a rapidly 
decreasing series (this condition which resem- 
bles that in M. obscura is apparently caused 
by abnormal reduction of the fourth supracil- 
iary); 26 rows of scales round midbody; 20 
lamellae under the fourth toe; five lower labials. 
Colour as for lectotype. Supranasal scales 
fused to nasals. If Storr is correct that sup- 
ranasals are invariably present in M. obscura, 
this specimen is conspecific with the lectotype. 
Paralectotype (c): This specimen is neither 
conspecific nor congeneric with the lectotype, 
and it differs in the following characters: 
snout-vent length 29 mm; tail (broken) 5 mm; 
length of forelimb 6 mm; length of hindlimb 
9 mm. Four fingers and five toes. Scalation 
greatly different: all supraciliaries fused into 
a single elongate shield; two supraoculars, the 
anterior very large and elongate, the posterior 
small; 18 scale rows round midbody; 16 lam- 
ellae under the fourth toe. Colour as for 
Menetia greyi Gray (see Boulenger 1887). 
This specimen is conspecific with the syntypes 
of Menetia greyi Gray (BMHN X.1.7. a-e; 
RR 1946.8.15.1-14 and 1946.8.16.88-99) 
and is referable to that species. 
Paralectotype (d) : Description as for para- 
lectotype (c) except: snout- vent length 27 mm; 
tail (broken) 3 mm; length of forelimb 6 mm; 
length of hindlimb 9 mm; 18 scale rows round 
midbody; 17 lamellae under the fourth toe. 
This specimen, is also conspecific with the 
syntypes of Menetia greyi Gray (see above) 
and is referable to that species. 
Type Series of Morethia anomalus Gray, 1845 



36 



P. A. RAWLINSON 



Remarks: Gray (1845) described this species 
from two specimens in the British Museum 
collection (BMNH) XI.6a-b; RR 1946.8.15. 
74-75). Starr (1972) designated one (BMNH 
XL 6b; RR 1946.8.15.75) as lectotype, thus 
making M. anomalus a junior subjective syn- 
onym of M. Uneoocellata, but did not comment 
on the identity of the other syntype. The author 
has re-examined Gray's two syntypes and 
determined that they belong to different spec- 
ies: the lectotype is conspecific with the lecto- 
type of M. Uneoocellata which is designated 
and described above; but the paralectotype is 
conspecific with Storr's (1972) M. obscura 
described at the same time that the lectotype 
of M. anomalus was designated. 
Lectotype: Storr (1972): BMNH XL 6b; RR 
1946.8.15.75, British Museum of Natural His- 
tory, London. Locality: West Australia. Col- 
lector: Mr. Gilbert. No other data. 
Description: Snout-vent length 43 mm. Tail 
(intact) 63 mm, 149% of snout-vent length. 
Length of forelimb, 11 mm; length of hind- 
limb, 17 mm. Snout-axilla 9 mm; axilla-groin, 

25 mm. Six supraciliaries, third, fourth and 
fifth equal in size and penetrating deeply be- 
tween the supraoculars; sixth supraciliary the 
smallest. Supranasal separate from nasal. Post- 
nasals present and separate from supranasal. 
One ear lobule. Eight upper labials, sixth the 
largest and entirely subocular; seven lower 
labials. One pair of nuchals. Eight preanal 
scales, four slightly enlarged. Smooth scales, 

26 rows round midbody. Subdigital lamellae 
smooth, 20 under the fourth toe. Palmar tuber- 
cles apically rounded. 

Colour: Very faded due to preservation. Dorsal 
surface light brown, ocellations not visible. 
Light dorsolateral stripe faintly visible. Black 
upper lateral band and white midlateral stripe 
faintly visible. Ventral surface unmarked, 
white. 

This specimen is conspecific with the lecto- 
type of M. Uneoocellata (MNHP 3092) des- 
cribed above. As Storr (1972) did not exam- 
ine the syntypes of M. Uneoocellata when he 
designated the lectotype of M. anomalus and 
placed it in the synonymy of M. Uneoocellata, 
his decision was apparently based only on the 



written description of the latter species. The 
author has examined Storr's lectotype of M. 
anomalus and found it to be conspecific with 
the lectotype of M. Uneoocellata so Storr's 
action is now verified. 

Paralectotype: BMNH XI.6a; RR 1946.8.15. 
74, British Museum of Natural History, Lon- 
don. Locality: West Australia. Collector: Mr 
Gilbert. No other data. 

Description: Snout-vent length 50 mm. Tail 
(broken) 7 mm. Length of forelimb, 12 mm; 
length of hindlimb, 19 mm. Snout-axilla, 15 
mm; axilla-groin 30 mm. Six supraciliaries, 
fourth the largest and fourth, fifth and sixth 
form a rapidly decreasing series; the third and 
fourth supraciliaries penetrate deeply between 
the supraoculars. Supranasals separate from 
nasals. Postnasals present and separate from 
supranasals. Three ear lobules. Seven upper 
labials, fifth largest and entirely subocular; six 
lower labials. One pair of nuchals. Eight pre- 
anal scales, four slightly enlarged. Smooth 
scales, 26 rows round midbody. Subdigital 
lamellae smooth, 18 under the fourth toe. Pal- 
mar tubercles apically rounded. 
Colour: Very faded due to preservation. Dorsal 
surface light olive-grey, ocellations not visible- 
Black upper lateral band faintly visible. Ven- 
tral surface unmarked, white. 

This specimen is not conspecific with the 
lectotype of M. Uneoocellata, but it is con- 
specific with Storr's species M. obscura (see 
below) and fits Storr's description of that 
species exactly. 

Diagnosis: Six supraciliaries (occasionally five 
owing to fusion of first and second), the third, 
fourth and fifth subequal and penetrate deep- 
ly between the supraoculars; the sixth supracil- 
iary the smallest. Supranasals usually fused to 
the nasals. Subdigital lamellae smooth or 
obtusely keeled. Palmar tubercles apically 
rounded. 

Description: (After Smyth (1972) and Storr 
(1972)). Snout-vent length 19-49 mm. Intact 
tail 111-247% of snout-vent length. Suprana- 
sals normally fused to nasals or separated only 
by a shallow or incomplete groove. Postnasals 
normally present but only separated from nasal 
by a faint groove. Frontonasal wider than 



THE ENDEMIC AUSTRALIAN LIZARD 



37 



long. Frontal longer than than wide. One to 
three ear lobules. Midbody scales in 24-31 
rows (usually 26 or 28), mean 27-3. Lamellae 
under the fourth toe 16-26, mean 19-7. 
Colour: Head coppery brown. Dorsal surface 
green, olive-grey or olive-brown, usually mark- 
ed with white ocelli outlined in black. Ocelli 
may be absent or modified into black or white 
spots which sometimes fuse into longitudinal 
stripes. White dorsolateral stripe present. Irreg- 
ular dark brown or black upper lateral band. 
White midlateral stripe margined by black 
below usually well developed, runs through 
ear, over forelimb and along trunk to hindlimb. 
Distribution: On the mainland restricted to 
two coastal areas in the SW of Western Aus- 
tralia: the mid-west coast from Port Cloates 
S to Geraldton; and the lower W coast from 
just N of Perth S to Cape Leeuwin and a 
short distance inland. Also occurs on islands 
of the Montebello Group and Houtmans Abrol- 
hos, and from Rottnest and Garden Islands. 
Distribution inland very limited. Not known 
from any other Australian state. (Figure 4.). 




Fig. 4. M. lineoocellato 



Literature Records: See list in Storr (1972). 
Specimens Examined: Western Australia (AM) 
Bunbury, R 30343: (QM) Rottnest Island, 
J 12241: 

Morethia obscura Storr 1972 

(Fig. 5.). 

Morethia anomalus (part) Gray, 1845, Catalogue of 
lizards: 65. 



Ablepharus Hneoocetlatus B var. anomalus (part) 

Boulenger, 1887, Catalogue oj the Lizards in the 

British Museum (Natural History) 3: 348-349. 
Ablepharus lineoocellatus (part) Zietz, 1920, Rec. 

S. A ust. Mas. 1: 220-221. 
Ablepharus lineoocellatus anomalus (part) l.overidge, 

1934, Bull. Mus. Comp. Zool. 11: 377-378. 
Morethia lineoocellata Smyth, 1972, Rec S. Aust. 

Mus. 16: 1-14. Figs. 4, 6. 
Morethia obscura Storr, 1972, J. R. Soc. W. Aust. 

55: 73-79. Figs. 1, 3. 

Holotype: WAM R 16916, Western Australian 
Museum, Perth. Locality: 6 miles cast of Kal- 
amunda, Western Australia, 31° 58' S, 116° 
08' E. Collector: Mr John Dell. Date of col- 
lection: November 7, 1962. 
Description: Sec Storr (1972). 
Remarks: M. obscura, described by Storr in 
1972, is very closely related to M. lineoocel- 
lata. Neither Smyth nor Storr examined the 
syntypes of M. lineoocellata though both used 
the name in their reviews. Smyth recorded 
and described specimens from South Australia 
under this name. However, he expressed reser- 
vations as he noted that South Australian 
specimens diflcrcd from many Western Aus- 
tralian specimens in several respects viz.: the 
invariable possession of supranasal scales; hav- 
ing the fifth supraciliary smaller than the 
fourth; and having only the third and fourth 
supraciliaries penetrate between the suprao- 
culars. Smyth noted that Gray (1845) des- 
cribed M. anomalus from 'W Australia' and 
distinguished it from M. lineoocellata because 
the former, but not the latter, had supranasal 
scales. Smyth did not use the name M. anom- 
alus for South Australian specimens as he 
doubted that the presence or absence of sup- 
ranasal scales on its own was a good indicat- 
ion of a species. He suggested that a careful 
study of Western Australian material was 
needed as both lineoocellata forms occurred 
there and stated that this could result in the 
recognition of two species. Storr carried out 
such a study and separated the new species 
M. obscura from M. lineoocellata using the 
condition of the supraciliaries described by 
Smyth and the presence of supranasal scales 
as the major diagnostic characters. All eastern 
Australian specimens, including those described 
by Smyth as M. lineoocellata, have proved to 
be M. obscura. 



38 



P. A. RAWLINSON 



The nomenclature of the species presently 
recognized in the genus Morethia has now 
become very confused, especially in the lineo- 
ocellata group. Literature records of M. lineo- 
ocellata prior to Smyth and Storr could include 
any, or all, of the 'southern' species of More- 
thia and should be disregarded unless they 
can be verified. Particularly confusing is the 
history of the species M. anomalus described 
by Gray (1845) from Western Australia. As 
mentioned above, Gray distinguished the spec- 
ies from A. lineoocellatus because it possessed 
supranasal scales. Boulenger (1887) after ex- 
amining material in the British Museum which 
included Gray's types of M. anomalus, applied 
the name Ablepharus lineoocellatus anomalus 
to specimens with supranasal scales. Authors 
from that time mainly followed Boulenger (e.g. 
Loveridge, 1934 and used the name anom- 
alus for the more eastern populations of M. 
lineoocellata which possess supranasal scales 
(i.e. Storr's M. obscura), though, as Smyth 
notes, most authors were probably also includ- 
ing M. adelaidensis and M. boulengeri under 
this name. As recorded above, there were two 
syntypes of Gray's M. anomalus, the lectotype 
(BMNH XI. 6b, RR 1946.8.15.75) which is 
conspecific with M. lineoocellata sensu stricto, 
and the paralectotype (BMNH XI.6a, RR 
1946.8.15.74) which is conspecific with Storr's 
new species M. obscura. The name M. anomala 
could have been retained if the paralectotype 
had been nominated instead as lectotype for 
the taxon. This would also have preserved 
recent usage of the name. It is unfortunate 
that the name anomalus has been placed in 
the synonymy of M. lineoocellata and a new 
name, M. obscura, has been introduced. It 
must be stressed again that in Smyth's review, 
the name M. lineoocellata was applied to speci- 
mens which now properly belong in Storr's 
species M. obscura and all Smyth's descriptions 
etc. apply to this latter species. 
Diagnosis: Six supraciliaries (rarely five owing 
to the fusion of the first and second), fourth 
the largest, and the fourth, fifth and sixth form 
a rapidly decreasing series, the third and fourth 
(and rarely the fifth) penetrate between the 
supraoculars, sixth the smallest. Supranasals 



invariably present. Subdigital lamellae smooth 
or obtusely keeled and unicarinate; palmar 
tubercules apically rounded. 
Description: Snout-vent length 1 8-56 mm, 
mean 43 mm. Total length of adults with intact 
tails 107-129 mm, mean 117 mm. Intact tail 
120-189% of snout-vent length. Supranasals 
and postnasals always present but often fused 
to each other or only separated by a shallow 
groove. Supranasals widely separated. In most 
specimens the third and fourth supraciliaries 
penetrate between the supraoculars, the fourth 
is largest, and the fourth, fifth and sixth are 
successively smaller; but rarely the fifth sup- 
raciliary is nearly as large as the fourth and 
it also penetrates between the supraoculars. 
Frontonasal wider than long. Frontal longer 
than wide. One pair of nuchals. One to four 
ear lobules. Midbody scales in 24-31 rows 
(usually 26 or 28), mean 27-7. Lamellae 
under the fourth toe 14-23, mean 190. 
Colour: Olive-brown to olive-grey dorsal sur- 
face, usually with dorsal ocellations which 
consist of a single scale with the middle third 
white and the outer thirds black. The dorsal 
ocellations are rarely bold or numerous, and 
may be reduced to black flecks or be absent 
altogether. Occasionally there is a trace of a 
pale dorsolateral stripe. Broad irregular black 
upper lateral stripe. Narrow pale irregular mid 
lateral stripe usually present, running from eye 
through ear over forelimb and back to hind- 
limb. The upper lateral and mid lateral stripes 
are not as even or bold as those in M. bou- 
lengeri. 

Distribution: Arid and semi-arid areas of SW 
New South Wales; NW Victoria; S South 
Australia and offshore islands; and S Western 
Australia. (Figure 5.). 

Literature Records: See lists in Smyth (1972 
as M. lineoocellata) and Storr (1972). 
Specimens Examined: Western Australia: 
(AM) Perth, R 2457; Western Australia, R 
6483; Bornham, R 7689; Cranbrook, R 7690; 
Merredin, R 9152; Eradu near Geraldton, R 
9164; Woodlands, Tambellup, R 11121; R 
11666; Northam, R 12350: South Australia: 
(NMV) Purnong, D 1546-51, D 3076; Lake 
Wangary, Eyre Peninsula, D 15058: New South 



THE ENDEMIC AUSTRALIAN LIZARD 



39 



Wales: (AM) Nymangee, R 17675; Round 
Hill Fauna Reserve between Lake Cargelligo 
and Mt. Hope, R 27860; R 27869; R 27823-4; 
R 27833; R 27838; R 27883-4; R 29670-1; 
8 km W of Nymangee, R 18481: Victoria: 




Fig. 5. M. obscura 



(NMV) Raak Plains D 698; Ouyen, D 1040; 
D 2416; Red Cliffs, D 7969; Kiata, Little 
Desert, D 8958; Pink Lakes near Ouyen, 
D 9473-4; Kooloonong, D 13951; Broughtons 
Waterhole, Little Desert, D 14917-29; li km 
NE of Broughtons Waterhole, D 14930-1; 
D 14937; 8 km S of Broughtons Waterhole, 
D 14932-3; D 14935; D 14942-3; D 14945; D 
14947-8; H km E of Broughtons Waterhole, D 
14934; D 14936; D 14940; D 14944; D 14946; 
H km W of Broughtons Waterhole, D 14938; 
5 km E of Broughtons Waterhole, D 14939; 
3 km NNE of Broughtons Waterhole, D 14941; 
Stan's Camp, Little Desert, 22 i km SW 
of Nhill, D 14949-50; 16 km N of Goroke, 
Little Desert, D 14951-2; 14i km S of Murray- 
ville, D 18235; Spring, 48 km S of Murrayville, 
D 18236; SA-Vic. border due W of Telopea 
Downs, Big Desert, D 38819; Red Bluff, Big 
Desert, D 38822; D 40174-5; The Springs, 
Big Desert, D 38831; 15 km E of Broughton, 
NW of Nhill, D 44866: 6 km S of Dimboola, 
D 44867: Little Billy Bore, Big Desert, D 
47392-3: No accurate data (AM) R 39455. 



Ecology 

Detailed laboratory studies have been made 
only on M. boulengeri. Field observations show 
M. adelaidensis and M. obscura to be similar 
to M. boulengeri and, from the distributions 
of M. butleri and M. lineoocellata, it is con- 
sidered that they would also be broadly similar. 

M. boulengeri is a heliothermic, insectivorous 
skink. The species has high thermal prefer- 
ences compared to other skinks from other 
temperate areas (Rawlinson, 1974 a,b; 1975). 
The voluntary minimum temperature is 
29-95°C, the mean preferred temperature is 
3409°C, and the voluntary maximum temp- 
erature is 39-35°C. 

Habitat preferences have not received any 
detailed attention. However, it is possible to 
state that all species live in open vegetation 
forms ranging from semi-desert to woodland. 
Morethia species are generally restricted to 
areas where mean annual rainfall is less than 
50 cm. 

Reproduction 

M. adelaidensis, M. boulengeri and M. obscura 
are all oviparous, but details are known only 
for M. boulengeri. Unlike the majority of 
lygosomid skinks from cool temperate areas, 
this species does not show obligatory sperm 
storage overwinter (Rawlinson, 1974 b). 
Ovarian and testicular activity commences in 
early spring (September to October) and 
ovulation occurs in late October to early Nov- 
ember. Copulation and fertilization also occurs 
at this time. The fertilized eggs are retained 
in the oviducts until late January or early 
February when they are laid in an advanced 
state of development. Clutch size varies from 
3 to 5 with a mean of 3-5 (11 observations). 

Relationships 

There is no doubt that the five southern 
species of Morethia are all closely allied and 
that with the northern (M. taeniopleura) group 
they form a good genus. The relationships of 
the genus have been discussed several times 
recently (see Greer 1974 and included authors). 
Greer recorded that the Morethia species have 
an 'alpha' lygosomid palatal bone pattern that 



40 



P. A. RAWLINSON 



fits them into his 'Group II* of Leiolopisma-like 
genera. On this basis he concluded that the 
closest Australian genera are Anotis, Crypt- 
oblepharus, Emoia, Leiolopisma and Pseu- 
demoia (though he included the latter genus 
in Leiolopisma) and the present author agrees 
fully with this finding. However, using morpho- 
logical criteria, Greer went on to construct 
a phylogeny in which the 'Group IF Leiolo- 
pisma-likQ genera were derived from an an- 
cestral species close to Pseudemoia spenceri 
(which he placed in the genus Leiolopisma). 
Greer considered the genus to be the end of 
an evolutionary line that ran from a Pseudem- 
oia spenceri-like. ancestor through Emoia to 
Cryptoblepharus then Morethia, In a second 
lineage Greer considered that the genera Anotis 
and Proablepharus arose from Leiolopisma via 
the same Pseudemoia spenceri-\\ke ancestor. 
The present author considers this phylogeny 
to be unlikely as it involves the evolution of 
five oviparous genera (Anotis, Cryptoblepharus 
Emoia, Morethia and Proablepharus) through 
two groups (Pseudemoia and Leiolopisma) that 
are placental, viviparous forms. Until full 
details of the biology and morphology of all 
'Group II' Leiolopisma-like species are known, 
their origins, relationships and phylogeny must 
remain doubtful. 

Discussion 

The five southern species of Morethia pre- 
sent an interesting pattern of speciation in the 
temperate arid and semi-arid areas of Australia. 
However, until further details of the ecology 
of the various species are known, it is not 
possible to meaningfully comment on their 
biogeographic and evolutionary significance. 
When these details are known, the group will 
provide a radiation pattern to compare and 
contrast with other related lygosomid groups, 
the more mesic temperate genera Lampropho- 
lis, Leiolopisma and Pseudemoia, and the 
tropical genera Anotis, Cryptoblepharus and 
Carlia. 

Acknowledgements 

The author thanks Dr H. G. Cogger of 
the Australian Museum, Mr A. J. Coventry 



of the National Museum of Victoria, and Ms 
J. Covacevich and Mr G. Ingram of the 
Queensland Museum, for help in locating 
specimens in the collections under their care. 
Ms A. Grandison, Dr E. Arnold and Mr A. 
Stimson of the British Museum of Natural 
History in London, Professor J. Guibe of the 
Museum National d'Historie Naturelle in Paris 
and Dr G. Peters of the Zoologisches Museum 
der Humboldt in Berlin all gave invaluable 
assistance in the location and examination of 
type specimens and associated data. Finally 
the author wishes to express sincere appreci- 
ation for the help given by various staff mem- 
bers of the National Museum of Victoria, espe- 
cially to Mr A. J. Coventry and Mr T. A. 
Darragh for assistance with the manuscript 
and Mr J. McNally for allowing the facilities 
of the Museum to be used during the collect- 
ion of data. 

References 

Boulenger, G. A., 1887, Catalogue of the Lizards in 
the British Museum (Natural History) 3: 3 18- 
349. 

Dumeril, A. M. C. and G. Bibron, 1839. Erpeto- 
logie Generale 5: 817. Paris. 

Fuhn, J. E., 1969. The 'polyphyletic' origin of the 
genus Ahlepharus (Reptilia, Scincidae) : a case of 
parallel evolution. Z. zool, syst. Evolutionsforsch 
7: 67-76. 

Gray, J. E., 1845. Catalogue of the specimens of 
lizards in the collection of the British Museum: 
65. British Museum, London. 

Greer, A. E., 1967. A new generic arrangement for 
some Australian scincid lizards. Brevoria No. 
267: 1-19. 

, 1970. A subfamilial classification of scincid 

lizards. Bull. Mus. Comp. Zool. Harv. 139: 
151-183. 

; — , 1974. The generic relationships of the 

scincid lizard genus Leiolopisma and its relatives, 
Aust. J. Zool. Suppl. Ser. No. 31: 1-67. 

Guibe, J., 1954. Catalogue des types des lezards. 
Colas, Bayeux O.P.I.A.C.L., Paris. 

Loveridge, A., 1934. Australian reptiles in the Mu- 
seum of Comparative Zoology, Cambridge, Mas- 
sachusetts. Bull. Mus. Comp. Zool, Harv. 77: 
377-378. 

Lucas, A. H. S. and C. Frost, 1895. Preliminary 
note of certain new species of lizards from Cen- 
tral Australia. Proc. R. Soc. Vict. 7: 264-269. 

Mittleman, M. B. 1952. A generic synopsis of the 
subfamily Lygosominae. Smithson. Misc. Collect 
117: 1-35. 

Ogilby, J. D., 1890. Redescription of an Ahlepharus 
from Australia. Rec. Aust. Mus. 1: 1-10. 

Peters, W., 1 874. Uber einige neue Reptilian 
(Lacerta, Eremias. Diploglossus, Euprepes, Lygo- 



THE ENDEMIC AUSTRALIAN LIZARD 



41 



soma, Sepsina, Ablepharus, Simotes, Onychoce- 
phalus). Sber. Dt. Akad. Wiss. Phys. — Math. 
Klasse. Juni 1874: 367. 

Rawlinson, P. A., 1974a. Biogeography and ecology 
of the reptiles of Tasmania and the Bass Strait 
area. Ch. 11. In Williams, W. D. (Ed.), Bio- 
geography and Ecology in Tasmania. Mono- 
graphic Biologicae 24: 230-269. The Hague, Junk. 

, 1974b. Revision of the endemic south- 
eastern Australian lizard genus Pseudemoia 
(Scincidae: Lygosominae). Mem. natn. Mus. 
Vict. 35: 87-96. 

-, 1975. Two new lizard species from the 



genus Leiolopisma (Scincidae: Lygosominae) in 



southeastern Australia and Tasmania. Mem. 

natn. Mus. Vict. 36: 1-15. 
Smyth, M., 1972. The genus Morethia (Lacertilia, 

Scincidae) in South Australia. Rec, S. Aust. Mus. 

16: 1-14. 
Storr, G. M., 1963. Ablepharus butleri, a new scincid 

lizard from Western Australia. W. Aust. Nat. 9: 

46-47. 
, 1972, The genus Morethia (Lacertilia, Scin- 
cidae) in Western Australia. /. R. Soc. West. Aust. 

55: 73-79. 
Zietz, F. R., 1920. Catalogue of Australian lizards. 

Rec. S. Aust. Mus. 1: 181-228. 



A CREEPING CTENOPHORAN (PLATYCTENEA: CTENOPHORA) 
FROM VICTORIA, AUSTRALIA 

By Brian J. Smith and Rhyllis J. Plant 
Invertebrate Department, National Museum of Victoria, Melbourne 

Summary 

auu f ? P ?o&J ens ° f 1 a P[ at y c , tei ? e an ctenophoran tentatively identified as Coeloptana wtlteyi, 
Abbott, 1907, are described living on red and green algae at the southern end of Port Phillip 
Bay, Victoria. These constitute the first record of this group of animals from southern Australia 
and only the second record for Australia, 



Introduction 

During survey work at the southern, end of 
Port Phillip Bay, Victoria, Mr Phil Hollis of 
the Underwater Research Group of Victoria 
discovered several specimens of a small creeping 
ctenophoran. These were discovered on detailed 
examination of minute algal faunas in an 
aquarium following collection of bottom 
growths from selected deep water areas in the 
Bay, and were brought in alive to the Museum 
where they were observed for several days. 
They were readily identified as platyctenean 
ctenophorans belonging to the genus Coelo- 
plana by their creeping habit, the presence of 
two retractile pinnate tentacles, a central dorsal 
statocyst and the absence of swimming comb- 
plates. 

This constitutes the first record for the Order 
Platyctenea for southern Australia and is only 
the second record for Australia, the other being 
for the Great Barrier Reef (Stephemson, 1931). 
Apart from an otherwise unpublished record 
by Dayton and Robillard for Antarctica 1968 
(pers. comm. in Gordon, 1969) it is the most 
southerly record for this group of unusual 
animals. 

MATERIAL 

Four specimens were discovered on algal 
growth taken from 15 5 m in a tidal hole 
1-5 km north of Portsea in the southern part 
of Port Phillip Bay, Victoria (38° 19' 5; 144° 
45' E). All specimens were found by Mr Phil 
Hollis, the first in December 1972 and three 
more in January 1973. The first and second 
specimens were found crawling on Caulerpa sp. 
while the others were on red algae. The samples 
in each case consisted largely of algae though 



some ascidians and other sessile animals were 
also present in small numbers. It is therefore 
not known with certainty on what substratum 
the animals were originally taken. However, 
they are not confined to one type of substratum 
and appear to be able to readily pass from 
one to the other. 

Three specimens are preserved in 5% neutral 
formalin in the National Museum of Victoria, 
Reg. No. G2649. 

DESCRIPTION 

The animal has a very flat flexible body 
capable of extension in any direction, with a 
thicker central dome-like region. In the centre 
of this domed region is a statocyst, composed 
of a central granule suspended in a vesicle, 
which is overlaid by a section of the body wall. 
There is an aperture in the outer body wall, 
which is figure-8 shaped with a central constric- 
tion, the long-axis of the aperture being per- 
pendicular to the inter-tentacular axis. This 
aperture can be opened and closed very rapidly, 
presumably by a sphincter muscle system. 
Surrounding the statocyst in the central area 
are several clear vesicles or pustules. On the 
few specimens examined these pustules vary in 
number from 8 to 25, ranging in size from 
three times as large as the vesicle containing 
the statocyst to approximately half its size, 
and in organization from a regular arrangement 
of radiating lines around the statocyst to a 
totally irregular arrangement. 

There are two long pinnate tentacles capable 
of extension to 6 to 8 times the body diameter. 
These tentacles are completely retractable into 
two tentacular sheaths at opposite sides of the 
body in the central domed region. When the 



43 



44 



BRIAN J. SMITH and RHYLLIS J. PLANT 



tentacles are retracted the positions of the 
tentacular sheaths are barely discernible as 
slightly raised, smaller areas of the central 
dome region. 

The overall body colour is a dark pink to 
orange-red with small pale or transparent areas 
and some white blotches. This appears to be 
made up of small red pigment spots and some 
white pigment spots in a largely transparent 
general body structure. Contraction of the body 
causes an intensification of the colour. 

The ventral surface is flat, with less colour, 
giving the body a semi-transparent appearance. 
The general distribution of the main body or- 
gans can be seen through the ventral surface. 
The position of the tentacular sheaths can be 
seen and also a series of canals surrounding a 
meridional canal. 

BEHAVIOUR 

When crawling the animals expanded to 12 
mm in diameter and the tentacles were capable 
of extending to at least six to eight times the 
body diameter. The animals were first ob- 
served crawling on the surface of green or 
red algae. Tentacles were streamed either to- 
gether or independently and retracted intermit- 
tently. The animal crawled freely over the 
surface of the alga with no particular part 
leading. However, several independent obser- 
vations were made of the body being bent 
round so that the two tentacular sheaths were 
positioned on the same side, allowing both 
tentacles to be streamed in the same direction. 

The animal was dislodged from the alga and 
was observed to swim feebly by a series of 
undulating wave movements of the thin peri- 
pheral region. It was also observed to gain the 
surface film of water in a shallow dish and to 
float inverted on the surface film by completely 
expanding its under-surface. While floating in 
this manner the tentacles were fully streamed 
several times. On one occasion the tentacles 
touched the bottom of the dish and appeared 
to adhere to it for a short period. During this 
time they were slowly contracted, pulling the 
animal along. 

When stimulated with a needle, when either 
crawling or floating, the tentacles were retracted 
very rapidly and completely. After a short 



period, either one or both were very slowly 
expanded in stages, each stage being inter- 
spersed with further rapid complete contrac- 
tions. No feeding activity was observed. 

IDENTIFICATION 

Platyctenean ctenophorans have been re- 
corded from many parts of the world, from 
Greenland (Mortensen, 1912) to Antarctica 
(P. Dayton and G. A. Robilliard 1968 in 
Gordon 1969), with records from practically 
every contient. However, they are still such 
rare and unusual animals that they have promp- 
ted study whenever they have been discoverd 
and many new taxa have resulted. This has 
been especially evident when a specimen has 
been discovered for the first time in a marine 
faunal zone or geographical region in which 
this group of animals has not previously been 
recorded. This phenomenon, coupled with the 
absence of any clearly defined taxonomically 
useful characters, has led to a proliferation of 
specific and generic names and taxonomic 
confusion. 

Several partial revisions of the group, have 
been undertaken, principally by Dawydoff 
(1936, 1938), Komai (1934) and others, 
while a good general account is provided by 
Hymen (1940). Following this latter work the 
present specimens are referred to the genus 
Coeloplana because of the presence of erectile 
dorsal papillae, the absence of comb-plates, and 
because the statocyst and tentacles are in hidden 
tentacular sheaths. The specimens are tenta- 
tively identified as Coeloplana willeyi Abbott 
(1907), following Matthews and Townsley 
(1964) and Gordon (1969), because the size, 
colour and number of dorsal papillae fall within 
the variation range of this species. However it 
is recorded as such, more to provide a con- 
venient label for future reference than with 
any idea of taxonomic exactitude. 

Acknowledgements 

We would like to thank Mr Phil Hollis for 
first discovering these animals and making 
them available for study. 

References 

Abbott, J. F., 1907. The morphology of Coeloplana. 
ZooL Jl. (Anat. Ontog.) 24: 41-70. 



CREEPING CTENOPHORAN 



45 






s: 



© 



Figs. 1, 2 and 3 — Series of drawings of Coeloplana 
willeyi Abbott of the living animal showing 
the general body shape and the tentacles. 

Fig. 4 — Central statocyst. 

Fig. 5 — Animal floating on surface film, using the 
tentacles for locomotion. 



46 



BRIAN J. SMITH and RHYLLIS J. PLANT 



Dawydoff, C, 1936. Les Ctenoplanidae des eaux de 
l'lndochine Francaise. Etude systematique. Bull, 
biol. Fr. Belg., 70: 456-486. 

, 1938. Les Coeloplanides Indochinaise. 

Arch. tool. exp. gen. 80: 125-162. 

Gordon, D. P., 1969. A Platyctenean Ctenophore 
from New Zealand. N.Z. 31. mar. Freshwat. Res., 
3: 466-471. 

Hyman, L. H., 1940. The Invertebrates. Part 1. Pro- 
tozoa through Ctenophora. McGraw-Hill, New 
York. 



Komai, T., 1934. On the structure of Ctenoplana. 
Mem. Coll. Sci. Kyoto Univ., Ser. B. 9: 245-256. 

Matthews, D. C. and S. J. Townsley, 1964. Addi- 
tional records of Hawaiian Platyctenea (Cteno- 
phora). Pacif. Set., 18: 349-451. 

Mortensen, T., 1912. Ctenophore I. Tjalfiella tris- 
toma Mrtas. Dan. Ingolf. Exped., 5(2): 3-59. 

Stephenson, T. A. et al., 1931. The structure and 
Ecology of Low Isles and other reefs. Sclent. 
Rep. Gt. Barrier Reef Exped., 3: 17-112. 



SYMBIOCLADIUS AURIFODINAE sp. nov. (DIPTERA, CHIRONOMIDAE), 
A PARASITE OF NYMPHS OF AUSTRALIAN LEPTOPHLEBIIDAE 

(EPHEMEROPTERA) 

By H. B. N. Hynes 

Department of Biology, University of Waterloo, Ontario, Canada 

Summary 

Symbiocladius aurifodinae sp. nov., an orthoclad chironomid parastic on nymphs of the 
mayfly genus Atalophlebioides, is described from mountain streams in Victoria. Descriptions 
are based on mature pupae and larvae collected during the summer. It is concluded that there 
is probably only one generation per year. This is the first record of Symbiocladius from 
Australia, and it is shown that this species is closely related to S. wygodzinskvi Roback from 
Argentina. 



Introduction 

Fontaine (1964) and Arvy and Peters 
(1973) have reviewed the literature on the 
larvae of Chironomidae that are found in asso- 
ciation with mayfly nymphs. It appears that 
the only species which actually feed on the 
tissues of their hosts are members of the genus 
Symbiocladius, and most records are from 
the flattened nympths of the Heptageniidae of 
the Northern Hemisphere. There is, however, 
one record of larvae on a species of Lepto- 
phlebiidae from North America (Mayo 1969), 
and careful description of another type of 
larva from a flattened leptophlebiid from sou- 
thern South America (Roback 1965). It may 
be noted here that flattened leptophlebiid 
nymphs replace the Heptageniidae in the 
Southern Hemisphere. 

The larvae found by Mayo (1969) on 
nymphs of Thraulodes were identified as Sym- 
biocladius, but it is clear from her description 
and figures that they differ considerably from 
the other described species. The larvae, for 
instance, seem not to be parasitic, they have 
well-formed head capsules and they retain 
eyespots, caudal bristles and anal gills. They 
seem, in fact, to be not unlike the phoretic 
genus Plecoteracoluthus, which occurs on per- 
lid stoneflies (Steffan 1965) and Megaloptera 
(Hilsenhoff 1968). 

The specimens from Argentina, on the other 
hand, are clearly parasitic and damaging to 
their hosts, which were tentatively identified 
as Thraulodes. It was intriguing therefore, 
especially in view of the similarity of the biotas 



of southern South America and Australia, to 
find a very close relative on nymphs of the 
leptophlebiid Atalophlebioides in streams on 
the Great Dividing Range in Victoria. 
THE MATERIAL 
The specimens were obtained during monthly 
collections (June 1971 to June 1972) of the 
fauna of several streams that were used in the 
study of the life histories of stoneflies. The 
methods used are described by Hynes and 
Hynes (1975), where more information on the 
streams is given. Mayfly nymphs carrying larvae 
or pupae of Chironomidae were found in only 
three of the 11 stream stations that were in- 
tensively studied. 
These were: 

Crown Creek above Woods Point (map re- 
ference 424367), 2300 ft, a cool (max. 
16i°C), swift stream 5-10 m wide and up 
to 40 cm deep, with a stable bed of rocks 
and shingle. 

Godfreys Creek below Frenchman's Gap 
(421374), 2500 ft, a cool (max. 13*°C), 
fairly swift shallow stream 2-3 m wide, with 
a stable stony and gravelly bed containing 
some silt. 

Delatite River below Sawmill Settlement 

(434423), 1900 ft, a cool (max. 16*°C), 

swift, turbulent river about 15 m wide and 

up to at least 1 m deep, with a stable bed 

of boulders, stones and coarse sand. 

No specimens of Atalophlebioides, which is 

a common genus in stony streams, were seen 

with chironomid larvae in any of the many 

other streams that we visited in Victoria. How- 



47 



48 



H. B. N. HYNES 



ever, one pupa was taken, and unfortunately 
lost in an attempt to breed it out, in Leather- 
barrel Creek, N.S.W. (615491) on January 8, 
1974. 

Two larvae were first noticed in the collec- 
tion made on October 26, 1971, in Delatite 
River, one was found on November 24 in 
Crown Creek, and several were found in God- 
freys Creek on December 28. They persisted 
in small numbers until February and March in 
the two creeks; in December there were many 
pupae, and in January and February only pupae 
(one pre-pupa in March) were obtained. In 
Delatite River only pupae were found on De- 
cember 28 and no specimens were collected on 
or after January 25. These findings possibly 
indicate only a single generation per season 
for the chironomid. It seems that at least some 
species of the host mayfly, which is present at 
all times, are uni voltine ( Duncan 1 972 ) . 
Thirty-seven larvae and pupae were collected 
in total, 20 from Godfreys Creek and 12 from 
Delatite River, and four mayflies carried empty 
shrouds from which, presumably, pupae had 
emerged. 

The Hosts 

The host mayflies appear to be all of the 
same species of Atalophlebioides (Figure 1), 
which it is possible may later become specifi- 
cally identifiable by its comb-like tarsal claws 
and by the peculiarly thickened distal margins 
of two of the segments near the bases of the 
three tails (Figures 1, C and D). No counts 
of uninfected specimens were made, but it is 
estimated that only about 1% carried the 
chironomid. 

Infected nymphs are all middle-sized, 3i to 
61 mm long, whereas fully developed nymphs 
were 8-8i mm long. There was good evidence 
that the chironomid caused stunting rather 
than that smaller nymphs were selected for 
infestation. The larvae were attached laterally 
to the thorax, indifferently as to side: 18 out 
of 41 were on the right. When wing pads were 
present the one nearest to the parasite was 
always reduced (Figures 1, A and B) and the 
entire development of the nymph seems to have 
been retarded. For instance, the uninfected 
nymph of which the mesonotum is shown in 



Figure 1, F, was the same size and, as judged 
by the developing male eyes, in the same instar 
(probably the penultimate) as the specimen 
shown in Figure 1, B. Also, the most fully 
developed infected nymph, a 6i mm male that 
still carried an empty pupal shroud, seemed to 
be physiologically near emergence in that its 
better developed wing pad enclosed a folded 
structure; but its wings were tiny and distorted 
(Figure 1, E) as compared with normal speci- 
mens in late instars (Figure 1, G and H). It 
may also be significant that the only chironomid 
collected on March 29 was a pre-pupa 1*75 
mm long, which is shorter than the smallest 
formed pupa that was collected, and it was 
taken on one of the two smallest infected 
mayflies (3i mm). It was also the only larva 
collected after the end of December. One may 
suppose that it infested a nymph that was too 
small or too unhealthy for ordinary develop- 
ment to occur. 

The parasite larvae were completely enclosed 
in a tough membranous shroud, as has been 
described for other species of Symbiocladius 
(Codreanu, 1939; Roback, 1965), and they 
lay alongside the nota, sometimes with the tail 
tucked under the wing pad as shown in Figure 
1, A. The larval head was directed forward 
or backward, apparently indifferently (13 out 
of 25 were forward), but I could find no 
trace of lesions on the nymphs. It seems clear, 
however, that, as with the European species 
(Codreanu, 1939), the host must supply the 
food. 

The pupae were all attached as shown in 
Figure 1, B, with their heads over the mayfly 
abdomen, and it appears from the empty 
shrouds that they leave by a dorsal rip. There 
was no relationship between the sex of the 
host and that of the pupal parasites; all four 
possible combinations were found among the 
12 pupae collected. 

THE PARASITE 

Six specimens, two larvae and four pupae, 
were used for the preparation of microscope 
slides for comparison with the clearly closely 
similar Argentinian species S. wygodzinskyi 
(Roback, 1965). The mountant was Euparal. 




LEPTOPHLEBIIDAE NYMPHS PARASITES 



49 





B 



Fig. 1 — A and B, Nymphs of Atalophlebioides with 
a larva and pupa of Symbiocladius aurifo- 
dinae (larger scale line 1 mm) : C and D, 
claw and base of filum terminale of Atalo- 
phlebioides (upper scale line 0-1 mm): E-H, 
mesonota of Atalophlebioides (larger scale 
line 1 mm), E, an infected specimen 6i mm 
long, F-H, uninfected specimens 6i, 7i and 
8i mm long. H, from Delatite River Decem- 
ber 28, 1971, E, Godfreys Cr. February 25, 
1972, rest from Godfreys Cr. December 28, 
1971. 



Genus Symbiocladius Kieffer 
Subgenus Acletius Roback 
1965 Entomol. News 76: 114-115. 

The Australian material agrees in most 
respects with Roback's definition of Acletius. 
These include the haired eyes of the adult 
(Figure 2, E and H), the subequal tibial spurs 







H 



(Figure 2, D), the long pectinate empodium, the 
three long basal spines on the claws (Figure 
2, G), the latero-dorsal position of the larvae 
on the host (Figure 1, A), the five lateral teeth 
on the labial plate (Figure 3, B and E) and 
the two teeth on the mandibles (Figure 3, C). 
There are, however, small differences. The 
antennae of the female have seven segments, 
rather than six (Figure 2, H), and the two 
mandibular teeth of the larva are subequal 
rather than being one robust and one accessory 
(Figure 3, C). I was unable to observe the 
palpal segments of the adult. 

Symbiocladius (Acletius) aurifodinae sp. nov. 

Male described from pupal material; 2-2 
mm. Head and thorax dark brown, abdomen 



50 



H. B. N. HYNES 










Fi £- 2 — Sytnbiocladius aurifodinae. A and B, D-G 
holotype, H and I, allotype. A, genitalia and 
part of pupal skin (smaller scale line 0-1 
mm): B, tip of dististyle (larger scale 01 
mm): C, ditto of specimen from Godfreys 
Cr December 28, 1971: D, tibial spurs of 
hind leg (smaller scale line 01 mm): E, 
optical section of eye margin (larger scale 
line 0-1 mm): F, pupal thoracic respiratory 
organ (smaller scale line 01 mm); G, claw 
of hmd leg (smaller scale line 001 mm): H 
and I, head and abdominal tip of female 
(smaller scale line 01 mm). 

brown with paler patches at bases of hairs. 
Antennae with at least 14 segments; eyes hairy, 
the hairs about as long as the diameter of the 
facets (Figure 2, E). Thorax apparently with 
bristles omly on scutellum. Pronotum narrow 
and collar-like. Legs with subequal tibial spurs 
and with claws with three basal hairs (Figure 2, 
D and G); segment ratios as in Table 1. 
Genitalia as Figure 2, A and B, but note that 
the number of bristles at the tip of the dististyle 
is three in the type and five in the other speci- 
men (Figure 2, C). 



TABLE 1 

Symbiocladius aurifodinae ratios of leg seg- 
ments of type specimens (100 = 07 mm) 



tarsal segments 
femur tibia 12 3 4 5 



Male 

fore 100 200 130 88 49 16 17 
mid 99 146 131 34 27 16 15 



hind 



134 163 108 60 33 21 21 



Female 
fore 



161 



225 171 64 37 29 29 



Female described from pupal material; 31 
mm. Similar to male, and also to S. wygod- 
zinskyi Roback (Figure 2, H and I). It has, 
however, seven antennal segments, no ventral 
hair on the antennal pedicel, and it lacks the 
paler spots at the bases of the abdominal 
hairs. 

Pupa, length, male 2-2 to 2-8 mm (holo- 
type 2-2), female 2-3 to 3-8 (allotype 31); 
small respiratory trumpets present (Figure 2, 



LEPTOPHLEBIIDAE NYMPHS PARASITES 



51 





B 







Fig. 3 — Symbiocladius aurifodinae, larval appendages 
(scale line 001 mm): A, antenna: B, labial 
plate: C. mandible: D, Labrum: E, labial 
plate and tip of labrum: F, maxillary palp: 
A-D, head capsule of holotype: E and F, 
head capsule from a female pupa from 
Delatite R. December 28, 1971. 



F) . Cuticle thin and pale brown, without spines. 
In the male a small featureless tail fin (Figure 
2, A); genital sacs elongate (Figure 2, A, 
where the tips of the sacs are missing). Note 
that Roback (1965) states that in S. wygod- 
zinskyi the anal fins are twice the length of 
the genital sacs, but this does not agree with 
his figure. In mature female pupae, in which 
the abdomen is distended with eggs, the anal 
fin is hardly apparent. 

Larva, the larvae range in length from 0-5 
to 2-1 mm (two that are certainly last instar, 
with developing appendages, are 2 and 2-1 
mm). Head pale yellow with a darkened hind 
edge; capsule widely open behind, almost a 
hemisphere. No eyespots. Body light brown 
with well formed anterior prolegs and small 
posterior ones; no posterior hairs or gills. An- 



tennae small and little sclerotized; number of 
segments uncertain (Figure 3, A). Labial plate 
broad with five lightly sclerotized teeth on each 
side (Figures 3, B and E). Maxilla shows only 
a small lightly sclerotized ring with several 
central conical structures (Figure 3, F). Man- 
dibles conspicuous hooks with two accessory 
teeth (Figure 3, C). Labrum with an apical 
swelling bearing about five lightly sclerotized 
teeth (Figures 3, D and E). 

MATERIAL EXAMINED 

Descriptions made mostly from the six 
mounted specimens. 

Holotype — mature male pupa, Crown Cr., 
Jan. 27, 1972. 

Allotype — mature female pupa, Godfreys Cr., 
Jan. 27, 1972. 

Paratypes — male pupa and 2 last instar larvae, 
same data as allotype. 
— head capsule of larva extracted 
from case of female pupa, Dela- 
tite R., Dec. 28, 1971. 



52 



H. B. N. HYNES 



Other Material 

— 2 larvae, 1 prepupa, 1 female pupa, 
Crown Cr., Nov. 24 and Dec. 28, 
1971, and March 29, 1972. 
— 11 larvae, 2 male pupae, 2 female 
pupae, Godfreys Cr., Dec. 28, 
1971, and Feb. 25, 1972. 
— 9 larvae, 3 female pupae, Delatite 
R., Oct. 26, Nov. 23 and Dec. 28, 
1971. 
All the specimens were collected by H. B. N. 
and M. E. Hynes. The types, paratypes and 
much of the other material has been presented 
to the National Museum of Victoria in Mel- 
bourne. Two larvae and one pupa from Delatite 
R., and four larvae and two pupae from God- 
freys Cr. remain in the author's collection. The 
species is named for the goldmines which are 
still active around Woods Point near to which 
most of the specimens were collected. 

Although in the absence of fully developed 
specimens, it is difficult to be certain, adults 
would probably run down to the genus Cric- 
topus in Freeman's (1961) key to the Aus- 
tralian Orthocladiinac. The larvae and pupae 
are, however, very different from those of CricCh 
topus. They are also quite distinct from those 
of Trissocladius, with which the genus Symbio- 
cladius has sometimes been combined, as was 
already pointed out by Saether (1969). 

S. aurifodinae is clearly very closely related 
to S. wygodzinskyi, but there are a few dif- 
ferences. The male has more antennal flagellar 
segments, at least 14 as opposed to 13, and the 
bristles at the dististyle tip are pointed not ovate. 
Th|C female has one more antennal segment. 
The pupa lacks dorsal spines and possesses 
a small respiratory organ. The larva has, ap- 
parently, much less robust lateral labial teeth 
and a wider central part to the labial plate, 
and its second mandibular tooth is larger. Also 
the tip of the labium which carries the spines 
is more swollen, and there seem to be no lateral 



spines below it such as are figured for S. wygo- 
dzinskyi by Roback. These seem to be very 
small changes after what must be many tens of 
millions of years of isolation of the two con- 
tinents. 

Acknowledgements 

I have previously acknowledged the help 
from many people and organizations that al- 
lowed me so rewarding a sabbatical year in 
Australia. My principal benefactors were the 
University of Waterloo, Monash University, 
and the National Research Council of Canada. 
I remain in their debt. 

References 

Arvy, L. and W. L. Peters, 1973. Phoresies, bio- 
coenoses et thnnatocoenoses chez les ephemerop- 
tcres. Proceedings of the First International Con- 
ference on Ephemeroptera 1970, Tallahassee, 
Florida, 244-312. 

Codreanu, R., 1939. Recherches biologiques sur un 
chironomid Symhiocladius rhithrogenac (Zavr.) 
ectoparasite 'cancerigene' des ephemeres torren- 
ticoles. Arch. ZooL exp. gen. 81: 1-283. 

Duncan, M. J., 1972. The life histories of Ephemerop- 
tera from two Victorian streams. B.Sc. Honours 
thesis. Department of Zoology, Monash Univer- 
sity. 

Fontaine, J. 1964. Commensalisme et parasitisme 
chez les larves d'ephemeropteres. Bull, mens. Soc. 
Lyon, 33: 163-174. 

Freeman, P., 1961. The Chironomidae (Diptera) of 
Australia. Aust. J. Zool. 9: 611-737. 

Hilsenhoff, W. L., 1968. Phoresy by Plecoptera- 
coluthus downesi on larvae of Nigronia serri- 
corm's. Ann. Entomol. Soc. Amer. 61: 1622-23. 

Hynes, H. B. N. and M. E. Hynes, 1975. The life 
histories of many of the stoneflies (Plecoptera) 
of southeastern mainland Australia. Aust. J. mar. 
Freshwat. Res. 26: 113-53. 

Mayo, V. K., 1969. Nymph of Thraulodes speciosus 
Traver with notes on a symbiotic chironomid. 
Pan-Pacif. Ent. 45: 103-112. 

Roback, S. S., 1965. A new subgenus and species of 
Svmbiocladius from South America. (Diptera: 
fendipedidae). Entomol. News 76: 113-122. 

Saether, O. A., 1969. Some nearctic Podonominae, 
Diamesinae, and Orthocladiinae (Diptera, Chiro- 
nomidae). Bull. Fish. Res. Bd. Can. 170: 1-154. 

Steffan, A. W., 1 965 . Plecopteracoluthus downesi 
gen. et sp. nov. (Diptera: Chironomidae), a 
species whose larvae live phoretically on larvae 
of Plecoptera. Canad. Ent. 97: 1323-44. 



THE ASCIDIAN FAUNA OF WESTERN PORT, VICTORIA, AND A 
COMPARISON WITH THAT OF PORT PHILLIP BAY 

By Patricia Kott 

Queensland Museum 
Abstract 

The taxonomy of 59 species of ascidians from Western Port and Port Phillip Bay, 
Victoria, is discussed. The ascidian fauna of Western Port is markedly more diverse than 
that of Port Phillip Bay. The biogeographical affinities of the species are assessed and the 
implications of the differences in species composition in the two areas are investigated. 



Introduction 

A previous collection of ascidians from Port 
Phillip Bay has been reported on by Millar 
(1966) but prior to that no major work has 
been devoted to the ascidian fauna of Victoria. 
The greater part of the present material has 
been collected for the National Museum of 
Victoria by the Underwater Research Group 
(Western Port Survey). Additional records of 
species occurring in Western Port, available 
from independent collections made by Mrs. J. 
Watson and Mr. K. Duke on parts of the 
adjacent Victorian coast, from Mallaooota 
near the Victorian — N.S.W. border to Portland 
Harbour and Cape Nelson, have been included 
in the present work. 

These collections are of particular interest 
in relation to the fauna of St. Vincent Gulf 
where large collections have recently been 
made and reported on (see Kott, 1972 a, b). 
Information on the better known fauna of Port 
Jackson and Moreton Bay to the north is also 
available. (Kott, 1952; 1957; 1962; 1963; 
1972 c, d). These locations are all large em- 
bayments in the Australian coastline, essen- 
tially marine, and tidal. They are, however, 
all protected from the direct swell of the 
southern ocean, and receive some fresh-water 
runoff from the water-ways emptying into them 
and from the shores surrounding them. 

The distribution of this sessile ascidian fauna 
is limited by the short free-swimming life of 
the pelagic larvae. Consequently species adap- 
ted to protected localities could be restricted 
in their distribution by lack of suitable sites 
for settlement on the open coast. The phylo- 
genetic relationships of the ascidian fauna of 



each of these sheltered embayments are there- 
fore of special zoogeographic and ecological 
interest in view of the likelihood of isolation 
of endemic and relict species. 

There are 59 species in the collections, of 
which one, Ciona intestinalis is probably in- 
troduced. These species, are set out in Tables 
1 and 2 together with others previously re- 
corded from Port Phillip Bay and Western 
Port but not represented in these collection. 
Aspects concerning the biogeography and habi- 
tat of the ascidian fauna of these locations is 
discussed below. 

Species List 

APLOUSOBRANCH1A 
CIONIDAE 

Ciona intestinalis 
CLAVELINIDAE 

Clavelininae 

Oxycorynia pseudobaudinensis n. sp. 

Podoclavella cylindrica 

Holozoinae 

Atapozoa mirabilis 

Sycozoa pedunculata 

Sycozoa cerebriformis 
POLYCITORIDAE 

Eudistoma pyriforme 
POLYCLINIDAE 

EUHERDMAN1INAE 

Pseudodistoma cereum 
Dumus arenijerus 

POLYCLININAE 

Polyclinum marsupiale 
Aplidium depressum 
Aplidium lobatum 
Aplidium triggiensis 



53 



54 



PATRICIA KOTT 



Synoicum hypurgon 

Synoicum sp.? 

Sidneyoides tamaramae 
DIDEMNJDAE 

Trididemnum cerebrijorme 

Trididemnum cyclops 

Didemnum candidum 

Didemnum spongioides 

Didemnum skeati 

Didemnum moseleyi 

Didemnum patulum 

Didemnum turritum 

Didemnum augusti 

Didemnum roberti 

Didemnum lambitum 

Lissoclinum fragile 

Lissoclinum ostrearium 

Diplosoma translucida 

Diplosma rayneri 

Polysyrwraton orbiculum 

Polysyncraton victoriensis n. sp. 
PHLEBOBRANCHIA 
ASCIDIIDAE 

ASCIDIINAE 

Phallusia depressiuscula 
Ascidia Sydney ensis 
Ascidia gemmata 
STOLIDOBRANCHIA 
STYELIDAE 

BOTRYLLINAE 

Botrylloides leachii 
Botrylloides nigrum 

POLYZOINAE 

Symplegma viride 
Amphicarpa diptycha 
Polyandrocarpa lapidosa 

STYEL1NAE 

Polycarpa thelypanes 
Cnemidocarpa etheridgii 
PYURIDAE 
Pyura australis 
Pyura cataphracta 
Pyura irregularis 
Pyura albanyensis 
Pyura lepidoderma 
Pyura scores biensis 
Pyura stolonifera praeputialis 
Halocynthia hispida 
Herdmania momus 



Microcosmus australis 

Microcosmus nichollsi 

Microcosmus helleri 

Microcosmus stolonifera 

Microcosmus squamiger 

MOLGULIDAE 

Molgula mollis 

Molgula sabulosa 

SYSTEMATICS 

Ciona intestinalis Linnaeus 

Ciona intestinalis Linneaus, 1767, p. 1087. Kott, 1952, 
p. 319 for synonymy and description. 

New Records: Port Phillip Bay (Oil wharf, 

Yarra River; artificial reef). 

Distribution: See Kott, 1952. 

Remarks: Kott (1969) has suggested that the 

cosmopolitan occurrence of this species, which 

is recorded from harbours and wharf piles in 

all regions outside the Antartic, is due to its 

transport on ships' hulls. 

Oxycotynia pseudobaudiensis sp. nov. 

(Fig. 1) 

Clavelina baudinensis Kott, 1957, p. 87 (part: speci- 
men with larger larvae) ?Millar, 1966, p. 363. Kott, 
1972a, p. 4. 

Type Locality: Laverton Bay (Victoria) 
Holotype: Australian Museum A.M. Y1113. 
Paratypes: W. Aust. Rottnest I., AM Y1112 
(Kott 1957). S. Aust.: Carickalinga Head, 
South Australian Museum S.A.M. E876; 
Rapid Head S.A.M. E 877 (Kott 1972a). 
Vict. (Western Port): Balnarring Beach, A.M. 
Y1122 (Kott 1957); Crawfish Rock; Flinders 
Jetty, N.M.V. (new records). 
Description: The colonies are 5-8 cm high 
and the wider terminal part of the head is 2 
cm in diameter terminally. In the upper half 
of the colonies the test is delicate and some- 
times glassy and transparent and encloses the 
body of the zooids which are never separate. 
On the upper surface the test forms only 
slight rounded protruberances over the anter- 
ior aspect of the zooids. In some colonies the 
zooid bearing upper part may be subdivided 
into several lobes. The slightly bulbous stalk 
narrows toward the base and is only slightly 
longer than half the height of the colony. The 
test of the stalk is firm and opaque and some- 
times slightly leathery externally. 



ASCIDIAN FAUNA OF WESTERN PORT 



55 



Zooids are from six to eight mm. In pre- 
servative they are a bluish colour and have 
dark accumulations of pigment in the mid line 
dorsal and ventral to the branchial 
siphon. There are 12 to 20 longitudinal 
muscles on the thorax and, depending on their 
degree of coalescence, they may vary in num- 
ber on each side of the body. From six to eight 
of the most ventral bands are aligned at a 
slight angle with the longitudinal axis of the 
body and break up into branches over the 
anterior half of the endostyle. Of the remain- 
ing longitudinal muscle bands more than half 
extend into the branchial siphon and the others 
into the atrial siphon. Posteriorly the bands 
extend along both sides of the abdomen. There 
are from 16 to 20 rows each of 20 to 30 
rectangular stigmata with a well developed 
transverse membrane between each row. In 
the mid dorsal line this membrane is expanded 
into the usual triangular, pointed languets. 

The gut forms a simple and fairly short loop 
(seldom longer than the thorax), enclosing 
the gonads. The anus opens at the base of the 
peribranchial cavity and is bordered with min- 
ute rounded lobes. The stomach has no true 
structural folds. Tt is present in the middle to 
posterior one third of the abdomen. There is 
no prestomach. 

Larvae are present only in the colonies from 
Rottnest Island and Laverton Bay (see Kott, 
1957: larger larvae). They are large, 0-9 mm 
long with the tail wound threequarters of the 
way around the body. Triradiate papillae are 
supported around a flattened frontal plate. 
The adhesive cells rise in a cone from tho 
centre of a depressed area which forms a 
fairly primitive papillary sucker or cup. The 
embryos appear to start their development in 
the proximal part of the oviduct and complete 
it in the right side of the peribranchial cavity 
where they demonstrate a wide range in stages 
of development. The most mature embryos 
are present anteriorly. 

Remarks: The separate identity of the present 
species was first suggested by the different 
larvae present in colonies from Rottnest and 
Victoria which had all been assigned to the 



species C. baudinensis Kott, 1952. Most of the 

previously described specimens of that species 
excepting only those recorded by Millar (1966) 
have been re-examined. 

Clavelina baudinensis from Cape Vlamingh, 
Rottnest Island and from Laverton Bay have 
small larvae (0-5 mm) in which the simple 
papillae without accessory suckers are sup- 
ported around the anterior end of the body 
which is not separated into a frontal plate. 
In these larvae the tail completely circles the 
body. These colonies can. be distinguished 
from the present species mainly by their long- 
er, narrow and cylindrical stalk. The anterior 
extremity of the zooids project more from the 
anterior surface of the test than in O. pseudo- 
baudinensis. Zooids of C. baudinensis exam- 
ined have a maximum of 12 longitudinal thor- 
acic muscles of which only a single band 
subdivides across the mid-line, ventral to the 
branchial aperture. In C. baudinensis there 
appears to be a more restricted range in the 
stage of development of embryos in the peri- 
branchial cavity. Although some eggs are 
present in the oviduct they do not appear to 
start their development there as in the genus 
Pycnodavella. In its colony and zooid form, 
C. baudinensis does appear to be closely re- 
lated to O. pseudobaudinensis; however, its 
larvae and the degree to which eggs are ap- 
parently fertilised in the atrial cavity suggest 
that it is a more primitive species. 

In O. pseudobaudinensis the oblique arrange- 
ment of the ventral thoracic muscles effects a 
depression of the anterior part of the thorax 
and draws it towards the postero-dorsal part 
of the thorax. The atrial aperture simultan- 
eously becomes terminal as in P. cylindrica, 
thus facilitating the liberation of large larvae. 
In C. baudinensis larvae are smaller and more 
easily liberated through the normally oriented 
aperture; and the more parallel arrangement of 
longitudinal thoracic muscles does not apocar 
to affect the relative position of the siphons. 
In neither of these species is the whole ventral 
surface withdrawn toward the postero-dorsal 
part of the thorax as is the case in Podocla- 
vella cylindrica where there is a special brood 
pouch ensuring the retention of embryos. 



56 



PATRICIA KOTT 



Podoclavella cylindrica (Quoy and Gaimard) 

Polyclinum cylindrica Quoy and Gaimard, 1834, p. 
618. 

Podoclavella cylindrica; Kott, 1972a, p. 5 and syno- 
nymy; 1972b, p. 167. 

New Records; Western Port (Flinders Jetty, 
Balnarring Beach). Ram Head (18 mis south 
of Mallacoota Inlet, 6 m). 
Distribution: W. Aust: Rottnest Island, Fre- 
mantle, Albany; S. Aust.: St. Vincent Gulf, 
West Island, Wright Island; Vict.: Bass Strait, 
Port Phillip Bay. The greatest recorded depth 
for the species is 22 metres (West Island, 
Kott, 1 972 ) . The species is known from 
sheltered caves and under ledges. 
Description: The present colonies generally 
have a firm basal common stalk and zooids 
are supported in the less firm terminal test 
and are independent of one another anteriorly. 
The usual blue pigment spots are present an- 
teriorly and thoracic musculature extends 
obliquely from the ventral border to the pos- 
tero-dorsal aspect of the body. Only in speci- 
mens from Western Port Bay are the zooids 
arranged around a central stalk which is 24 
cm long and 2-4 cm in diameter, thickest 
terminally where it breaks into branches. 
Larvae of the usual form without ampullary 
lobes, are present in the brood pouches of this 
colony. 

Remarks: In P. cylindrica contraction of the 
oblique thoracic musculature causes the fore- 
shortening of the dorsal length of the thorax by 
drawing the postero-dorsal corner ventrally and 
anteriorly. The branchial aperture is simultan- 
eously withdrawn leaving the atrial aperture 
terminal and anal and gonadial openings ad- 
jacent to it. The developing embryos are re- 
tained however, in a pouch from the dorsal 
surface, thus avoiding early liberation which 
could otherwise result from contraction of the 
thorax. 

The relationship of specimens in which 
there is a central stalk to those that are sup- 
ported upright and parallel to one another on 
a basal membrane is not known. Another 
specimen of the former type in the collection 
of the National Museum of Victoria is 60 cm 
long and resembles Distaplia cylindrica from 



the Antarctic (see Kott, 1969). The zooids 
in both forms are similar in every respect and 
the colonies appear to represent the same 
species. It is possible that the long axial 
stemmed forms are from deeper water. 

Atapazoa mirabilis Kott 
(Fig. 2) 

Atapazoa mirabilis Kott, 1972b, p. 168. 
New Records; Western Port (Tankerton jetty) 
Distribution: The species has previously been 
recorded from S. Aust. (Elliston Bay). 
Description: A single colony only is available. 
It is massive and irregular, 14 cm long and 
6 cm wide. It is composed of fairly thin layers 
of zooid bearing test that coalesce so that the 
colony is traversed by spaces. The atrial siphon 
from the postero-dorsal corner of the thorax 
is characteristically long and posteriorly direc- 
ted. Both the branchial and atrial apertures 
are bordered by six distinct lobes. There are 
three rows each of about 12 stigmata in each 
row. The horizontal gut-loop consists of a 
fairly long oesophagus, rounded stomach and 
wide intestinal loop. There is a single large 
ovum attached to the zooid from a region in 
the loop of the gut. The position of the brood 
pouch is apparently abdominal, rather than 
thoracic, and is reminescent of the situation in 
the Didemnidae. 

Remarks: The species has been recorded pre- 
viously only from Elliston Bay in South 
Australia. It is possible that it is endemic to 
the southern coast although the type location, 
on the floor of a cave, is only accessible to 
collectors equipped with SCUBA. 

Sycozoa pedunculata (Quoy and Gaimard) 

Aplidie pedunculatum Quoy and Gaimard, 1834, p. 
626. 

Sycozoa penduculata; Kott, 1972c, p. 234 and syno- 
nymy. 

New Records: Western Port (Rutherford 
Channel); Port Phillip Bay (no location, arti- 
ficial reef). South-east Portland. 
Distribution: W. Aust.: Cockburn Sound, King 
Georges Sound; Tas.: d'Entrecastaeux Chan- 
nel, Derwent Estuary, Furneaux Group; S. 
Aust.: St. Vincent Gulf; Vict.: Western Port, 



ASCIDIAN FAUNA OF WESTERN PORT 



51 



Port Phillip Bay, Lakes Entrance; Qd. : More- 
ton Bay. 

Discription: The usual large inverted conical 
heads on long slender stalks with basal tufts 
of roots. There are deep V-shaped furrows 
between each double row of zooids and the 
branchial apertures from each row of zooids 
open into each side wall of these furrows. The 
branchial openings are thus protected to some 
extent and the furrow provides an immediate 
microenvironment outside the openings. The 
apex of a rounded ridge between two of these 
furrows lies over the common cloacal canal. 
There are very large common cloacal openings 
around the outside of the flattened free ends 
of the lobes. 

Larvae are present in colonies from South- 
east Portland. They have the usual anterior 
papillae, an otolith, but no ocellus, and a 
short broad tail extending only three quarters 
of the ways around the body of the larva. 

Sycozoa cerebriformis (Quoy and Gaimard) 

Aplidie cerebriforma Quoy and Gaimard, 1834, p. 

625. 
Sycozoa cerebriformis; Kott, 1972a, p. 8 and syno- 
nymy. 

New Records: Western Port (Crawfish Rock, 
Tankerton Jetty). Portland Harbour (5-10 in 
on rocks forming jetty). 

Distribution: N.W. Aust; S. Aust.: St. Vincent 
Gulf; Vict. : Balnarring Beach (Western 
Port), Port Phillip Bay; N.S.W.: Jervis Bay, 
Port Stephens, Port Hacking, Port Jackson. 
South Africa. It has been recorded from 
5-40 m. 

Description: Specimens are sturdy fan-shaped 
colonies with short stalks. The fan or zooid 
bearing portion may extend into a thick 
undulating lamellae. Common cloacal aper- 
tures are present along either side of the flat- 
tened free edge of the lamella. The double rows 
of zooids converge from this outer edge of the 
lamella down toward the top of the stalk. 
Remarks: The specimens from deeper water 
at Crawfish Rock are larger than the colony 
taken in shallower water. It has already been 
observed (Kott, 1972a) that the species fav- 
ours areas where there are steady but not 
strong unidirectional currents, and no surge 



or turbulence. The fan-like colony shape is 
apparently adapted to take maximum advan- 
tage of this type of environment. This species 
appears to be confined to more sheltered 
locations than S. pedunculata, very often where 
there is some turbidity and a muddy substrate 
although in the absence of a larval ocellus 
the light conditions are not likely to affect its 
settlement. 

It has been recorded from a wider circum- 
polar range than S\ pedunculata but although 
it has been taken from north western Australia 
it is not recorded from Moreton Bay and it 
has not been recorded from Tasmania. Its 
latitudinal range is therefore more limited. It 
has not been taken from depths greater than 
40 metres. Its distribution suggests that it 
may represent a relict species confined within 
embayments in relatively shallow water where 
it can best take advantage of sheltered con- 
ditions. 

Polycitor giganteum (Herdman) 

Polyclinum giganteum Herdman, 1899, p. 79. 
Polycitor giganteum; Kott, 1972a, p. 9 and synonymy. 

New Records: Western Port (Crawfish Rock, 
Tankerton Jetty). Ram Head (18 mis south 
of Mallacoota Inlet, 6 m). 
Distribution: A wide circum-Australian dis- 
tribution from Rottnest Island (W.A.) and 
across the southern coast to Port Jackson (N. 
S. W.). 

Description: One specimen from Crawfish 
Rock is more or less flattened and sessile, 
about 13 cm in diameter but only 5 cm high; 
another specimen has the usual rounded gel- 
atinous head narrowing to a waist before ex- 
panding into a wide sandy base, possibly 
embedded in the substrate. The test is char- 
acteristically transparent and gelatinous and 
the zooids are large, radiating out from the 
base of the colony to open by separate 
apertures around the head. 
Remarks: The species appears to favour rocky 
substrates where -a firm adhesion can be 
effected, thus satisfying the requirements of 
a large, inflexible colony that is fixed by only 
a small area of the base. The species is found 
equally in embayments and on the open coast. 



58 



PATRICIA KOTT 



Eudistoma pyriforme (Hcrdman) 

(Fig. 3) 

Psammaplidium pyriforme Herdman, 1886, p. 419. 
Eudistoma pyriforme; Kott, 1972a, p. 9 and synonymy. 

New Records: Western Port (Crawfish Rock). 
Ram Head (18 mis. south of Mallacoota Inlet, 
6 m). 

Distribution: Palao and Gilbert Islands. Qd.: 
The Great Barrier Reef, in the Pacific; S. 
Aust.: St. Vincent Gulf. Madagascar. 
Description: The colony from Mallacoota Inlet 
is flattened and firm with sand absent only 
from the surface layer of test which has 
brownish-purple spherical pigment cells. The 
colony from Crawfish Rock is irregular and 
investing, with a dense sand inclusion through- 
out the test, making it rather hard and ob- 
scuring the arrangement of the zooids. There 
are about 10 fairly wide muscle bands down 
either side of the thorax and an almost con- 
tinous layer of circular muscles. The circular 
muscles on the siphons are well developed but 
do not form definite sphincters. The oesopha- 
gus is of medium length opening into a large 
shield shaped and smooth stomach halfway 
down the abdomen; the duodenal area is long, 
and in a contracted abdomen is bent in an 
S-shape. The intestine bends anteriorly after 
leaving a spherical posterior stomach, and 
forms a loop opposite the duodenum when the 
abdomen is contracted. The rectum extends 
anteriorly, to the peribranchial cavity, and is 
straight. 

Remarks: Although Hastings (1931) regarded 
the loop in the gut as diagnostic of this 
species it has been observed in other species 
where the abdomen is contracted. There are few 
reliable diagnostic characters available in this 
genus where colony shape is variable, no 
systems are formed, and where the strong 
body musculature results in highly contractile 
zooids. The development of the musculature, 
the length of the oesophagus and the nature 
of the common test, therefore, provide the 
only morphological characters to determine 
the species, and it is possible that some mis- 
identification occasionally occurs. The record- 
ed distribution of this species also suggests 



that more than a single species is represented. 

Pseudodistoma cercum Michaelsen 

Pseudod'tstoina cereurn Michaelsen, 1924, p. 364. Kott, 
1972a, p. 12 and synonymy. 

New Records: Western Port (Crawfish Rock). 
Cape Nelson (near Portland, vertical faces and 
roof of cave, moderate surge, 5 m). 
Distribution: The species is apparently com- 
mon intertidally and from depths down to 70 
metres off the South Island of New Zealand. 
Other records are from the eastern coast of 
Victoria, and off the South Australian coast 
and from Dakar. 

Description: The colony is very damaged and 
its form is not discernible. The test is very 
soft, jelly-like and transparent. Both apertures 
are 6-lobed, there are 3 rows each of about 
20 stigmata. Stomach folds are not apparent 
externally, however internally its glandular wall 
is interrupted in 4 places to give, the appear- 
ance of folds. The zooids are short and the 
thorax, abdomen and posterior abdomen are 
of equal length. 

Remarks: Zooids are characteristic. In view of 
the Australian and New Zealand records sug- 
gesting a circum-polar distribution in the south- 
ern cold temperate region the record from 
Dakar (Monniot, 1969) is surprising. Mon- 
niot's specimens do, however, agree with the 
present colonies and with those from New 
Zealand. The species is delicate and is taken 
from underneath ledges and in rocky locations 
where some protection is available. 

Dumus areniferus Brewin 
(Figs. 4, 5, 6) 
Dumus areniferus Brewin, 1952, p. 453. 
New Records: Western Port Bay (Crawfish 
Rock). 

Distribution: New Zealand: Otago. 
Description: The colonies form a thicket of 
elongate branching stalks, club-shaped termin- 
ally with the free end obliquely flattened. The 
outer test is encrusted with a single layer of 
sand particles giving rigidity to the otherwise 
extremely delicate test. Each terminal lobe con- 
tains only a single zooid. The maximum length 



ASCIDIAN FAUNA OF WESTERN PORT 



59 




Oxycorynia pseudobaudinensis 

1 contracted zooid, musculature not shown on 

abdomen. 
Eudistoma pyriforme 
2 — contracted zooid, musculature not shown on 

abdomen. 
Atapozoa mirabilis 
3 — zooid. 
Dumus areniferus 
4 — colony. 5 — zooid. 6 — larva. 



Aplidium depressum 
7 — zooid. 
Aplidium lobatum 
8 — zooid. 

Aplidium triggiensis 
9 — zooid. 

Synoicum hypurgon 
10 — zooid 

Sidenioides tamaramae 
1 1 — zooid. 1 2 — larva. 



60 



PATRICIA KOTT 



of the stalks is 60 cm. Zooids are about 3 
cm long, of which the long thread-like poster- 
ior abdomen is about two-thirds of the total 
length. Both apertures are 6-lobed and open 
directly onto the surface of the terminal flat 
surface of the stalks. There are no protective 
flaps of test protecting the apertures such as 
are found in Euherdmania australis. There are 
about eight very fine thoracic muscles along 
each side of the thorax. There are six long 
stigmata crossed by parastigmatic vessels in 
each of four rows. The abdomen is approxi- 
mately the same length as the thorax. The 
oesophagus is of moderate length, the smooth 
walled stomach is elongate and there is a 
posterior stomach and a duodenal region. A 
mid-intestinal region occupies, with the 
stomach and oesophagus, the descending limb 
of the gut loop before it enters the rectum in 
the pole of the loop. Testis follicles are pre- 
sent in a single row in the posterior abdomen 
and there is a group of ova anterior to the 
testis lobes, a little distant from the posterior 
end of the abdomen. The gonads occupy only 
the posterior half of the posterior abdomen. 

The larvae are 0-5 mm long and there may 
be up to nine in the peribranchial cavity. They 
have paired rows of ampullary vesicles dor- 
sally along cither side of the endostyle and 
along either side of the postero-ventral aspect 
of the larval body. There are the usual three 
papillae anteriorly and these alternate with 
median ampulllae. Lateral ampullae are also 
present either side of the median ampullae. 
Remarks: The specimens conform exactly both 
in colony and zooid form with those described 
by Brewin from New Zealand. The species 
resembles both Euherdmania australis and 
Ritterella herdmania, both with a similar col- 
ony. Externally it is distinguished from E. 
australis by the absence of the flap of test 
which protects the external aperture in the 
latter species (see Kott, 1957, 1972b), and 
from E, herdmania by the fan shape of the 
terminal tip of each lobe. It is possible that 
the aberrant colonies mentioned by Kott 
(1957) that were taken with E % australis were 
actually specimens of the present species. 



Polyclinum marsupiale Kott 
Polyclinum marsupiale Kott, 1963, p. 83. 
New Records; Western Port (Crawfish Rock). 
Distribution: S. Aust.: Victor Harbour; Tas.: 
Hunter Island; Qld. : Great Barrier Reef 
(Heron Island). 

Description: Colonies are mushroom-shaped, 
up to 2 inches in diameter across the upper 
surface of the head which is supported on a 
short stalk; or alternatively the colonies may 
be spherical and sessile, fixed by a small 
area of the base. There is a dense outer layer 
of sand on the test absent sometimes from 
parts of the upper surface. Internally the test 
is very soft with only very occasional sand 
grains included. The internal test is traversed 
by canals in which the zooids are contained, 
and forms only thin septa between these can- 
als. Preserved colonies are therefore often col- 
lapsed and flattened. 

The zooids, opening around the upper half 
of the colony, are very small. The branchial 
aperture is terminal. The antero-dorsal atrial 
aperture is on a short siphon with a circular 
sphincter muscle and the opening is protected 
by a pointed muscular languet from the body 
wall anterior to the siphon. There are 14 to 
15 rows of 10 to 12 rectangular stigmata with 
the usual rounded papillae on the transverse 
vessels. The stomach is smooth externally with 
an inner glandular wall. 

Remarks: The heads of living colonies are 
apparently distinctively spherical, although in 
preserved specimens the soft internal test col- 
lapses and they are flattened and sometimes 
appear lobed or folded. 

Aplidium depressum Sluiter 
(Fig. 7) 

Aplidium depressum Sluiter, 1909, p. 102. Kott, 1963, 
p. 95 and synonymy. 

New Records: Western Port (Crawfish Rock, 
Rutherford Channel) 

Distribution: Previously recorded only from 
Bargara (Queensland) and from Indonesia 
and the Philippines. The species is common in 
those locations from which it has been re- 
corded. The reason for these isolated records 
is not known. 



ASCID1AN FAUNA OF WESTERN PORT 



61 



Description; Soft, jelly-like, flat investing col- 
onies that are minute and circular fixed by a 
small area of their base, or more extensive 
fixed by the whole extent of the basal surface. 
The species commonly invests stalks and 
fronds of weed. Only very sparse sand grains 
are enclosed in the semitransparent brownish 
common test through which the zooids are 
clearly evident. In the smaller colonies the 
zooids are arranged in two or three circular 
systems of about six zooids, but in the larger 
colonies these expand into double row systems. 
The zooids are minute with an inconspicuous 
sessile atrial aperture halfway down the dorsal 
surface of the thorax. There is a single short, 
pointed atrial languet from the upper border 
of the aperture. There are five rows of about 
eight stigmata. The thorax and abdomen are 
of equal length and together represent half 
the length of the zooid. The stomach has 11 
distinct folds. 

Remarks: The small number of rows of stig- 
mata, with the number of stomach folds, the 
form of the colony and the nature of the test 
distinguish the species. 

Apldium lobatum Savigny 
(Fig. 8) 

Aplidittm lobatum Savigny, 1816, p. 182. Kott, 1963, 

p. 97 and synonymy. 
Non Psammaplidium lobatum; Herdman, 1899, p. 85 

«ApUdium solidum Herdman, 1899; Millar 1963; 

>A. arboratum Kott, 1963). 

New Records: Western Port (Crawfish Rock). 
Distribution: Florida, West Indies, the Mediter- 
ranean, Red Sea, Indonesia, Queensland and 
the Great Barrier Reef and New South Wales. 
The present record extends the southern range 
of this species from the east coast of Australia. 
Description: The colonies are irregular and 
investing. The common test is firm and hard 
with sand throughout. Zooids are minute with 
the thorax, abdomen and posterior abdomen 
all of equal length. The sessile atrial aperture 
has a deeply divided trifid languet from the 
anterior border of the opening. There are 8 
fine longitudinal muscle bands along each side 
of the thorax extending onto the abdomen and 
posterior abdomen. There are six rows each of 



about six stigmata. The four stomach folds 
are only apparent internally. 
Remarks: The species appears to be adapted 
for a rigorous environment, and is found in- 
vesting the undersurface of rocks and in the 
present case, in algal holdfasts. The firm test 
and zooids with few stomach folds, are similar 
to the condition found in A, solidum Herdman 
in which zooids open on both sides of flat 
lamellae. 

Aplidium triggiensis Kott 
(Fig. 9) 

Aplidium triggiensis Kott, 1963, p. 104. 
New Records: Western Port (Crawfish Rock). 
Distribution: W. Aust.: Rottnest Island, Triggs 
Island and Nornalup; Vict.: Balnarring Beach. 
Description: The colonies are very soft and 
investing stones, etc. Sometimes they are pro- 
duced basally into projections which extend 
into or around the substrate to form a very 
firm adhesion. Varying quantities of sand are 
present in the colonies. Posterior abdomina 
cross one another in the basal test. The zooids 
are minute, the thorax is 1-3 mm long and 
generally shorter than the abdomen when con- 
tracted; the posterior abdomen is long and 
thin and up to twice the length of the rest of 
the body. 

There is a sessile atrial aperture about one 
third of the distance down the dorsal surface 
of the thorax, with a short, pointed, undivided 
languet from the upper border of the opening. 
There are 9 to 10 rows of about 15 stigmata. 
The oesophagus is long and the stomach, pre- 
sent about half way down the abdomen, is 
broken up into 14 to 15 distinct folds. A single 
embryo is present in the peribranchial cavity. 
It is 0-6 mm long and anteriorly there is a 
multiplicity of adhesive papillae in the median 
line around the anterior end of the larva as 
previously described for this species (Kott, 
1963). 

Remarks: This species also appears to be 
adapted for very rigorous conditions, both by 
the form of the colony and its tendency to 
produce extensions to fix it firmly to the sub- 
strate. The larval form is quite distinctive and 
is large in relation to the size of the zooid. 



62 



PATRICIA KOTT 



Consequently, a maximum of two larvae have 
been reported as present in the peribronchial 
cavity. In the absence of this distinctive larval 
form, these species could be confused with 
Aplidium multiplicatum (Sluiter) which has 
been recorded from Queensland, Japan, the 
Philippines, Indonesia and from Broome, 
North-western Australia (see Millar, 1963). In 
the latter species, however, the posterior ab- 
domen is relatively short and the testis lobes 
form bunches in the posterior abdomen, rather 
than double rows, as in A. triggiensis. 

Aplidium pliciferum (Redikorzev) 

Atnaroucium pliciferum Redikorzev, 1927, p. 390. 
Kott, 1972a, p. 13 and synonymy. 

New Record: Western Port (Tankerton Jetty). 
Distribution: See Kott, 1972a. 
Description: The colony is a firm gelatinous 
cushion with a flat upper surface. There is 
a short stalk from the middle of the under 
surface. The margin of the colony is rounded. 
The test is semi-transparent and there is neither 
encrusting nor included sand. The zooids are 
tightly packed in double rows radiating from 
common cloacal apertures randomly placed 
on the upper surface. Anteriorly the zooids 
are parallel to one another and vertical, 
although the posterior abdomina may be more 
irregularly orientated in the basal half of the 
test. There are 12 fine longitudinal thoracic 
muscles extending separately along the abdo- 
men and both sides of the posterior abdomen. 
They are never gathered into a close band. 
The atrial aperture is sometimes produced into 
a short siphon and the pointed single bifid lip 
extends from the upper border of the aperture. 
There are 11 — 15 rows each of about eight 
stigmata. The thorax and abdomen are about 
the same length and the posterior abdomen 
is long and threadlike. The stomach has 18 
to 20 regular longitudinal folds. 

Larvae are present in the peribranchial cav- 
ity. They have a double row of ampullary 
vesicles from the lateral ridges on either side 
of the median papillae which alternate with 
median ampullae. 

Remarks: Although there is some variation in 
the number of rows of stigmata, in the number 



of stomach folds and in the shape of the 
colony, the firm gelatinous flat-surfaced colony 
form varies only in relation to the area by 
which it is fixed. The larvae are also character- 
istic. The species differs from the closely re- 
lated A, flavolineatum which has more thoracic 
muscles, more stomach folds, lateral branches 
on the larval median ampullae and no larval 
ampullary vesicles. 

Synoicium hypurgon (Michaelsen) 

(Fig. 10) 

Macroclinum hypurgon Michaelsen, 1924, p. 401. 
Synovium hypurgon; Kott, 1963, p. 86 and synonymy. 

New Records: Western Port (Crawfish Rock). 
Distribution: W. Aust.: Rottnest Island, Fre- 
mantle; Great Barrier Reef: Heron Island. 
New Zealand: North Island. 
Description: The present specimen consists of 
three large clavate to mushroom shaped lobes 
joined basally and to varying extends along 
their sides to form a large, hemispherical 
colony, 5 cm in diameter and sessile basally. 
There is sand around the sides of each lobe 
but not on the upper surface where the zooids 
open. The test is soft and gelatinous and has 
no foreign bodies. The zooids are present in 
the outer layer of the upper surface. There is 
a small, sessile atrial aperture one third of the 
distance down the dorsal surface of the thorax 
with a large, triangular atrial languet rising 
from the body wall anterior to the aperture. 
There are eight longitudinal muscle bands on 
the thorax. The branchial sac is very long 
and narrow with 13 rows of eight small oval 
stigmata in each row. The gut loop is short, 
about half the length of the thorax. The 
oesophagus is especially short and the stomach 
small and smooth. There is a duodenal en- 
largement and a posterior stomach in the loop 
of the gut. 

Remarks: Although there is considerable vari- 
ation in the form of colonies of this species 
and some variation in the amount of sand and 
other material which is contained in the com- 
mon test, the small zooids, long, narrow 
branchial sac and relatively short abdomen, 
together with the relative position of the atrial 
tongue from the body wall rather than from 



ASC1DIAN FAUNA OF WESTERN PORT 



63 



the anterior border of the atrial aperture dis- 
tinguish it. 

Synoicium sp.? 
Record: Western Port (Crawfish Rock). 
Description: The specimen is damaged and 
torn, although the fragments appear to re- 
present a fairly thin investing colony. The 
test is semi-transparent and very soft. Zooids 
appear to be arranged parallel to one another 
and vertical to the upper surface. Zooids are 
fairly small and the thorax and posterior ab- 
domen are about equal in length, while the 
abdomen is shorter. There are about 10 long- 
itudinal thoracic muscles. The atrial aperture 
is sessile and there is a single pointed languet 
from the upper margin of the opening. There 
are three rows of about 10 long rectangular 
stigmata, each row crossed by parastigmatic 
vessels. Dorsal languets are present in the 
mid-dorsal line opposite both transverse ves- 
sels and parastigmatic vessels. The stomach is 
shield-shaped and smooth without any areol- 
ations, although it has a glandular appearance. 
Remarks: The parastigmatic vessels in the 
branchial sac are unusual, although they have 
previously been described for Synoicium 
atopogaster Kott, 1962. The small number of 
rows of stigmata in the branchial sac suggests 
a relationship with Synoicium bowerbanki, 
which has, however, a longer oesophagus and 
a distinct atrial siphon. Further, in the present 
specimen, the dorsal languets opposite the 
parastigmatic vessels as well as the primary 
transverse vessels suggests that the rows of 
stigmata are in the process of subdividing and 
in fact the most posterior row does contain 
a few stigmata which are bisected in the region 
of the parastigmatic vessel. Synoicium papil- 
liferum differs from the present specimen in 
the presence of a long siphon, although it has 
a short oesophagus, as well as the same 
number of longitudinal muscles and about the 
same number of stigmata in each row, as 
does the present specimen. It is most probable, 
therefore, that this represents a juvenile of 
some species of Synoicium, rather than a new 
species characterised by 3 rows of stigmata 
crossed by parastigmatic vessels. 



Sidneioides tamaramae Kesteven 

(Figs. 11, 12) 

Sidneioides tamaramae Kesteven, 1909, p. 277. Kott, 
1957, p. 104. 

New Records: Western Port (Crawfish Rock). 
Distribution: N.S.W.: Tamaramae Bay. 
Description: The colonies are soft and pillar- 
like lobes. The free end of each lobe is raised 
into a rounded marginal ridge surrounding a 
terminal depressed surface from the centre of 
which there is a protruberant common cloacal 
aperture. The branchial apertures are made 
conspicuous by the absence of sand, around 
them. They open onto rounded swellings on 
the marginal ridge. The external test is com- 
pletely encrusted with sand, absent only from 
the region around the apertures. The test is 
otherwise very soft. The abdomen is about 
half the size of the long thorax. The atrial lip 
is narrow and fleshy but very long with about 
10 fine muscles extending along its length. The 
longitudinal thoracic muscles extend along the 
ventral side of the abdomen and the dorsum 
of the posterior abdomen causing it to curve 
when the muscles are contracted. There are 
17 rows of stigmata with 18 stigmata in each 
row. There is no sign of papillae on the 
transverse vessels. The stomach is oval with 
mulberry-like glandular swellings in its wall. 
There is a duodenal region, a posterior stomach 
and a mid-intestine which expands into the 
rectum before it curves into the ascending 
limb of the gut loop. The ovary is developed 
in the thoracic wall at about mid-thoracic 
level and projects into the peribranchial cavity 
just to the right of the mid-line, the vas 
deferens and the distal part of the rectum. The 
anal opening is opposite the 7th row of stig- 
mata. 

There are about 18 eggs at varying stages 
of development in the ovary. Free eggs are 
also present in the peribranchial cavity together 
with up to 20 developing embryos. Mature 
embryos are 6 mm long and the tail is 
wound completely around the bodv. There is 
a double row of vesicles along either side of 
the mid-dorsal line and a cluster of vesicles 
postero-ventrally on each side of the body. 
Paired lateral ampullae alternate with the three 



64 



PATRICIA KOTT 



anterior papillae but there arc no median 
ampullae. 

Remarks: This record has extended the range 
of this interesting species, previously regarded 
as endemic to a small region on the coast of 
New South Wales. 

I rididemnuin cyclops Michaclscn 

(Fig. 13} 

Trididannum cyclops Miehaelscn, [921, p. 19. Kott, 
L966, p. 286 and synonymy. Eldredge, 1967: 183. 

New Records: Western Port (Flinders Jetty, 
Eagle Rock). 

Distribution: West Indian Ocean. N. Aust. : 
Darwin, Great Barrier Reef. 
Description: Both the present records represent 
extensive colonies, almost completely investing 
specimens of Ascidia sydneyensis. h\ both 
cases the branchial aperture is free, although 
in one specimen the didemnid has grown over 
the atrial siphon, leaving a small space be- 
tween the test of the host through which the 
excurrent water could flow. 

The surface of the colony is smooth with 
a superficial layer of flat bladder cells and 
some spherical purple pigment cells. The spi- 
cules are dense beneath the layer of bladder 
cells and in the thoracic region and become 
less dense toward the base of the colony where 
they are absent altogether. They are from 003 
— 005 mm in diameter with up to 12 pointed 
rays in optical section. There are no zooxan- 
thellae in the common cloacal system of these 
specimens. There is a very shallow thoracic 
common cloacal cavity. The zooids 
have a minute thorax with three rows of 
stigmata. There is no endostylar pigment cap 
present in these specimens. The retractor 
muscle is fairly long. There is no atrial siphon, 
although there is a well defined and fairly long 
anterior lip from the border of the aperture. 
There is a single undivided testis follicle with 
8 J coils of the vas deferens. 
Remarks: The surface bladder cell layer, the 
shallow thoracic common cloacal canal, the 
absence of an atrial siphon and the form and 
distribution of the spicules are characteristic 
of this species. The extensive colonies are 



different to the typical small colonies of tropi- 
cal specimens. The absence of zooxanthellae 
should be especially noted as these are invari- 
ably present in specimens previously described. 
It is possible that the zooxanthellae are associ- 
ated with a tropical environment and should 
not be regarded as a specific character. In 
addition, the endostylar pigment cap has in- 
variably been present in previously described 
specimens of this species, but it is possible 
that its presence is a variable character as in 
7'. cerehrijorme (see below). However, these 
specimens do diverge from the characteristic 
facies of this tropical species and it is possible 
that there is a cline in its characters that is 
evident at the southern limit of its range. 

Trididcmnum cerebriforme Hartmeyer 

(Fig. 14) 

Trididcmnum cerehrijorme Hartmeyer. 1913, p. 139. 
Kott, J972d, p. 247; 1972e, p. 47 and synonymy. 
non i rhthlcnuutm cebrijorme\ Kott, 1972b, p. 178. 

New Records: Western Port (Crawfish Rock). 
Distribution: South and West Africa; Indian 
Ocean; S.W. Aust.; S. Aust.; Vict: Phillip 
Island; N.S.W.; Qd.; Gulf of Carpentaria. It 
therefore has a wide distribution in the south- 
ern temperate to subtropical regions and is 
absent only from the eastern Pacific and the 
Western Atlantic. 

Description: The colony is irregularly lobed. 
Branchial openings are conspicuous and slightly 
protruberant, owing to the density of spicules 
rilling the branchial lobes. There is the usual 
posteriorly directed atrial siphon. There is an 
extensive posterior abdominal cloacal system 
formed by canals traversing a central core of 
test. The spicules are less dense than at the 
surface. They are large, from 04 to 007 mm 
in diameter with five conical rays in optical 
section. The endostylar pigment cap is absent. 
Remarks: With the exception of the endostylar 
pigment cap the zooids and colony of this 
specimen are identical with those previously 
described. The pigment cap is also absent from 
the specimens of T. cyclops from this locality. 
Specimens from South Australia (Kott, 1972b) 
without a posteriorly directed atrial siphon are 
incorrectly assigned to this species. 



ASCIDIAN FAUNA OF WESTERN PORT 



65 



?Didemnum candidum Savigny 

(Figs. 15, 16) 

Didemnum candidum Savigny, 1816, p. 194. Kott, 
1972a, p. 19 and synonymy; 1972b, p. 179. 

New Records: Western Port (Crawfish Rock). 
Distribution: Cosmopolitan (see Kott, 1972a). 
Discription: Flat small pinkish colonies, the 
colour being due to the fairly sparse distri- 
bution of spicules allowing the zooids to show 
through. Spicules are mostly in the surface 
and basal test. There is an extensive thoracic 
common cloacal system. Zooids are about one 
mm long with four rows of stigmata. The 
anterior border of the atrial aperture is pro- 
duced into an atrial lip, forked terminally. 
There is a single testis follicle with 4i coils 
of the vas deferens. Spicules range from 02 
to 05 mm in diameter with conical pointed 
to needle-like rays. 

Remarks: The present young colonies have the 
spicules typical of this species together with 
the extensive thoracic common cloacal cavity. 
Zooids of more typical colonies are brown. 
The production of the anterior border of the 
atrial aperture into a lip is another character 
not usual for the species. 

Didemnum moseleyi (Herdman) 

(Fig. 17) 

Leptoclinium moseleyi Herdman, 1886, p. 272. 
Didemnum moselyeyi, Kott, 1972a, p. 19 and syno- 
nymy; 1972b, p. 179; 1972d, p. 249. 

New Records: Western Port (Crawfish Rock, 
Eagle Rock). 

Distribution: Pacific and Indo-Malayan region; 
circum- Australian. 

Description: Common, although not very num- 
erous. White investing colonies and small 
circular colonies on weed are available. Spi- 
cules are dense throughout the test which is 
rather brittle. The common cloacal canals 
are thoracic and zooids are enveloped in an 
independent thoracic sheath where the thorax 
crosses the common cloacal cavity. The sur- 
face layer of test is very thin indeed. In some 
colonies there are primary canals extending to 
abdominal level and surrounding discrete 
clumps of zooids which are embedded abdom- 
inally in the common basal test, although their 



thoraces are separate, each in a discrete thor- 
acic sheath. The surface layer of test in these 
colonies is depressed over the deep primary 
cloacal canals giving a cauliflower appearance 
to the surface of the colony. Large common 
cloacal openings are distributed randomly over 
the surface and some spicule filled papillae 
are also present on parts of the colony. The 
spicules are 001 to 003 mm in diameter 
with about seven pointed rays in optical sec- 
tion. The common cloacal system in this 
species does not appear to develop by proli- 
feration of double row systems but by a de- 
velopment of the primary cavity to envelop the 
thorax of each zooid as it is added to the 
system. Zooids are minute and colourless in 
formalin. Small spherical lateral organs are 
present either side of the thorax opposite the 
most posterior row of stigmata. There is a 
long retractor muscle. The vas deferens coils 
9i times around the undivided testis follicle. 
In one specimen large spherical vesicular 
cells are present in the surface test surrounding 
each branchial opening to form wide inter- 
secting circles that interrupt the dense spicules 
so that the surface appears to be pitted rather 
than smooth. The spicules, cloacal system and 
zooids otherwise conform with D. moseleyi. 
Remarks: The vas deferns in these colonies 
has more coils than is usual for the species 
although a wide range has been reported. The 
distribution and form of the rather constant 
spicules and the size and position of the 
lateral organs have been used to determine 
this species and to distinguish it from Didem- 
num candidum (see Kott, 1972a) which has 
a similar cloacal system and a similarly wide 
range recorded for the spirals of the vas def- 
erens. 

Didemnum patulum (Herdman) 
(Fig. 18) 

Leptoclinum patulum Herdman, 1899, p. 92. 
Didemnum patulum Kott, 1972a, p. 18. 

New Records: Western Port (Crawfish Rock, 
on Ecklonia holdfasts and investing other as- 
cidians; Eagle Rock). 

Distribution: N.S.W.: Port Jackson; S. Aust: 
St. Vincent Gulf. 



66 



PATRICIA KOTT 



Description: Colonies form large sheets. The 
surface is smooth. Smaller colonies may be an 
even, greyish colour, but larger colonies always 
have grey-blue to black mottled markings. 
There is a surface layer of bladder cells and 
beneath these numerous stellate pigment cells 
are distributed amongst the spicules to form 
the mottled markings that characterise the 
species. Spicules gradually become less dense 
toward the base of the colony. The pigment 
cells are especially concentrated in the test 
overlying the cloacal canal; occasionally they 
may extend into the test beneath the surface 
layer and beneath the common cloacal cavities 
and when this occurs the colony is almost 
black in colour. The spicules are stellate, with 
pointed or rounded rays. The majority of 
spicules are 03 - 004 mm in diameter with 
about seven rays in cross section. There are, 
however, less common spicules of similar form 
but larger diameter, up to 005 mm. There 
are also smaller spicules with up to 12 sharply 
pointed rays in cross-section similar to those 
found in Didemnum candidum. 

The zooids are embedded in the rather solid 
common test and open on both sides of 
cloacal canals. Primary cloacal canals some- 
times extend the whole length of the zooid 
and may extend slightly posterior abdominally. 
The secondary cloacal canals remain at the 
level of the thoraces. The surface test is thick 
and there is a long branchial opening. The 
upper border of the atrial opening is some- 
times produced into a lip. There are four rows 
of about eight stigmata. The basal layer of test 
in which the abdomina of the zooids are 
embedded is rather thick and gelatinous. 
Remarks: This species is the most common 
ascidian in the particularly rich ascidian fauna 
at Crawfish Rock. The characteristic marking 
caused by greyish-black stellate pigment cells 
overlying the common cloacal canals and the 
thick layer of basal test distinguish the species 
from Didemnum candidum in which there is 
the same variety in form of the spicules. The 
common cloacal system is also distinctive in 
that the thoraces of zooids are not completely 
enveloped bv the cloacal cavity as in D. can- 
didum and D. moseleyi but remain embedded 



in common test opening into the cloacal canals 
from their dorsal surface. Although the species 
has been recorded from Port Jackson and from 
St. Vincent Gulf it is never present in the same 
high density as at Crawfish Rock and it has 
never been reported from Port Phillip Bay. 

Didemnum turritum Michaelsen 
(Figs. 19, 20) 

Didemnum turritum Michaelsen, 1930, p. 521. Kott, 
1962, p. 319. 

New Records: Western Port (Crawfish Rock; 
Eagle Rock). 

Distribution: S. W. Aust.; S. Aust.: St. Vincent 
Gulf. 

Description: Pinkish investing colonies. Large 
cloacal apertures are scattered over the surface. 
The branchial apertures are also conspicuous 
owing to the density of spicules in the test 
covering the branchial lobes. A single lobe 
of the branchial aperture sometimes developes 
a hollow pointed papilla from its base. Small 
rounded pigment cells line the common cloa- 
cal cavities. The surface of the test may be 
depressed over the deep primary common 
cloacal canals to form furrows on the surface. 
Clumps of zooids are surrounded by these 
deep primary canals which sometimes extend 
posterior to the abdomina of zooids. The 
secondary cloacal cavities are thoracic. The 
spicules are regularly stellate with about seven 
conical rays in section and are 0-3 to 0-4 
mm in diameter. The thorax of each zooid 
is about 10 mm long with large oval lateral 
organs which occupy a pronounced pit in the 
thoracic wall along most of its length. The 
branchial siphon has a well defined circular 
sphincter muscle but is not very long although 
the surface layer of test is thicker than that 
of either D. candidum or D. moseleyi. The 
atrial aperture is extensive exposing most of 
the dorsal part of the branchial sac and some- 
times its anterior border is produced into a 
pronounced forked lip. The thoracic retractor 
muscle was not detected. 

There are four rows of about eight stigmata. 
The gonads were not distinguishable. 
Remarks: The species is readily recognized by 
the hollow pointed papillae associated with 



ASCIDIAN FAUNA OF WESTERN PORT 



67 




Trididemnum cyclops 
13 — spicules. 

Tridemnum cerebriforme 
14 — spicules. 
Didemnum candidum 
1 5 — zooid . 1 6 — spicules. 
Didemnum moseleyi 
17 — spicules. 
Didemnum patulum 
18 — spicules. 
Didemnum turritum 
19 — thorax. 20 — spicules. 



Didemnum augusti 

2 1 — thorax. 22 — spicules. 

Didemnum roberti 

23 — spicules. 

Didemnum spongioides 

24 — spicules. 25 — larva. 

Lissoclinum fragile 

26 — spicules. 

Diplosoma translucidum 

27 — zooid. 28 — diagrammatic cross section through 

colony. 
Polysyncraton victoriensis 
29 — spicules. 



68 



PATRICIA KOTT 



one of the branchial lobes of each aperture 
in certain limited areas. These papillae super- 
ficially resemble those sometimes occuring in 
D. moseleyi although in the latter species 
they are not hollow and are not specifically 
associated with the apertures. The thicker 
surface test, the relatively large zooid and the 
large oval lateral organs also distinguish the 
species. 

The hollow papillae protecting the branchial 
apertures are reminiscent of those in D. nek- 
ozita Tokioka, 1967, from the Palau Islands 
and the Philippines. The latter species, how- 
ever, has distinctive spicules and a thoracic 
cloacal system. 

Didemnum augusti Michaelsen 

(Figs. 21, 22) 

Didemnum augusti Michaelsen 1920, p. 39. Kott, 

1962, p. 323; 1972d, p. 247. 
1 Didemnum partiturn Tokioka, 1953, p. 191. 

New Records: Western Port (Crawfish Rock). 
Ram Head (18 miles south of Mallacoota 
Inlet). 

Distribution: S.W. Aust.; S. Aust.: Reevesby 
Island; Vict: Balnarring Beach; West Indian 
Ocean. 

Description: Very extensive, thin, investing 
colonies with dense white spicules, less dense 
only in the basal test. The spicules are stellate 
from 03 to 05 mm with 5 to 7 conical 
pointed rays in optical cross section. The 
surface of the test is furrowed and has a 
cauliflower-like appearance where the surface 
test is depressed over the deep primary can- 
als. The primary canals extend the whole length 
of the zooids between pillars of common test 
in which the abdomina are embedded. Only 
the dorsal aspect of the thorax is exposed to 
the common cloacal canals. Some secondary 
canals are present but the thorax is never 
enclosed in its own discrete sheath of test. The 
test along either side of the atrial opening is 
thickened but there is no lateral organ. The 
thorax is small, 0-6 mm, with four rows of 
stigmata. There is a retractor muscle always 
present. 

Remarks: The species is distinguished from 
D. turritum and D. moseleyi, which often have 



the same surface furrows and deep primary 
canals, by the solid pillars of test in which 
the zooids are embedded, by the very small 
thorax, the absence of a distinct lateral organ 
and by the large spicules with few conical rays. 

Didemnum roberti Michaelsen 
(Fig. 23) 

Didemnum roberti Michaelsen, 1930, p. 516. 
Didemnum ternatanum; Kott, 1972b, p. 179. 

New Records: Western Port (Crawfish Rock; 
Eagle Rock). 

Distribution: W. Aust.: Shark Bay; S. Aust: 
Elliston Bay. 

Description: Investing colonies with a smooth 
surface and dense spicules in the surface and 
basal layers of test. The common cloacal aper- 
tures have their borders stiffened with spicules 
and are very conspicuous. Some colonies are 
flattened but in some, rounded lobes are 
developed by a thickening of the basal test 
to form a central core of test. There are ex- 
tensive posterior-abdominal cloacal spaces and 
the zooids are suspended in clumps between 
the surface and basal layers of test, anchored 
basally by a short single strand of test, and 
at the surface by the branchial lobes of re- 
spective zooids. Secondary common cloacal 
cavities surround the thoraces of the zooid, 
each surrounded by a discrete layer of test 
and with a large lateral organ occupying most 
of each side of the thorax (as in D. turritum). 
The spicules are not so thick in the test 
surrounding the zooids. The surface test is 
fairly thick, again resembling D. turritum. 
Spicules are almost spherical, 002 to 004 
mm in diameter with rounded rays. The zooids 
are small. The testis follicle is undivided and 
has 7i coils of the vas deferens around it. 
Remarks: The present colonies diverge from 
Michaelsen's (1930) specimens only in the 
presence of spicules throughout basal or axial 
test. In the Shark Bay material the basal 
layer of spicules was confined to a layer 
beneath the posterior abdominal canals to 
form a sort of endoskeleton. 

Colonies from Elliston Bay (Kott, 1972b) 
which are identical with the present colonies 
from Western Port, were erroneously assigned 



ASCIDIAN FAUNA OF WESTERN PORT 



69 



to the species Didemnum ternatanum Gotts- 
chaldt. Although the three-dimensional com- 
mon cloaca and the size and form of the 
spicules are similar to those of D. ternatanum, 
the present species is distinguished from it 
by its external, oval lateral organ, by the more 
densely distributed spicules; by the multiplicity 
of common cloacal apertures and extensive 
colony; and by its firmer consistency. D. rob- 
erti is distinguished from D. bistratum Michael- 
sen, 1920 from West Africa by the form of 
its spicules (those of the latter species are 
spherical and hollow) and by its external 
lateral organ. D. spongioides also has a simil- 
arly labyrinthine common cloaca, but its spi- 
cules are stellate, with fewer, conical rays, and 
fewer coils of the vas deferens. 

Didemnum roberti has previously been 
described as yellow, or yellowish— no inform- 
ation is available on the in vivo colour of 
the present colonies. 

Didemnum spongioides Sluiter 

(Figs. 24, 25) 

Didemnum spongioides Sluiter, 1909, p. 67. Kott, 
1962, p. 318; Eldredge, 1967, p. 193. 

New Records: Western Port (Crawfish Rock). 

Distribution: Caroline Is.; Indonesia; W. Aust.: 

Rottnest Island; Tas.: Oyster Bay. The records 

suggest a circum-Australian distribution. 

Description: Colonies are rounded to conical 

with a terminal common cloacal cavity. The 

test is firm. Spicules are present in a layer 

beneath a surface layer of bladder cells at the 

level of the branchial siphons. They are less 

dense beneath this layer and are entirely 

absent from the test core that occupies the 

centre of the colony. The surface of the test 

is covered with minute spicule-filled pointed 

papillae that project through the bladder cell 

layer and, when magnified, give to the surface 

a spotted appearance. 

The spicules are stellate with about seven 

conical rays in cross section, and range from 

002 to 006 mm in diameter. An extensive 

common cloacal cavity separates the outer 

spicule and zooid bearing layer of test from 

the inner spicule free test core in which 

embryos develop. Cloacal canals extend into 



the zooid bearing layer but these do not 
separate clumps of zooids from one another. 
The openings of the common cloacal canals 
into the posterior abdominal chamber are 
shown by Sluiter (1909, Plate 6, fig. 9) and 
the ridges and trabeculae he describes are 
formed by the roof of the cloacal chamber 
enclosing abdomina of zooids projecting into 
the chamber, between the openings of the 
canals. These ridges and trabecula are not 
the imprint of the substrate on the base of 
the colony as Eldredge (1967) has suggested. 
Zooids are small. The thorax, when contracted 
is only 0-5 mm long. There is a wide atrial 
opening and a small rounded lateral organ 
opposite the 4th row of stigmata. The vas 
deferens coils 6i times around the undivided 
testis follicle. 

Embryos are packed in the central test core 
at the base of the common cloacal chamber, 
into which they move through the occasional 
strands of test that connect the surface layer 
to the central core. They are 0-9 mm long 
when mature, have an ocellus and an otolith, 
and four pairs of lateral ampullae. The tail 
winds once around the embryo. 
Remarks: The species is related to D. lambitum 
in the form of cloacal system and the spicules 
and is distinguished from that species by the 
presence of a bladder cell layer and by the 
larva in which the ampullae are not sub- 
divided. 

D. spongioides; Eldredge, 1967, differs from 
the present specimens in the presence of pig- 
ment cells, the even investing form of the 
colony and in the condition of the cloacal 
system with well developed thoracic secondary 
canals and primary canals extending postero- 
abdominally but not forming a continuous 
space separating surface from central or basal 
test. The thickness of the surface layer of 
test, the arrangement of cloacal canals, the 
spicule form, size and arrangement, and the 
presence of pigment cells of Eldredge's speci- 
mens are identical with those of D. turritum 
from which they differ only by the absence 
of hollow pointed papillae on the surface. 

The colony is typically "sponge-like" in 
external appearance, rounded and sessile. 



70 



PATRICIA KOTT 



Didemnum lambitum (Sluiter) 

Didemnoides lambitum Sluiter, 1900, p. 18. 
Didemnum lambitum; Kott, 1962, p. 317 and syno- 
nymy; 1971, p. 19; 1972a, p. 18. 

New Record: Port Phillip Bay (Hobson's Bay). 
Distribution; N. Z.: Chatham Island, North 
Island, South Island, Stewart Island; Tas.; S. 
Aust. ; St. Vincent Gulf. 
Description: The colonies are more or less 
fan shaped, made up of vertical lamellae or 
columns. These may fuse for the greater part 
of their length, or only basally or terminally. 
The free outer edge of the fan is more or less 
flattened. Common cloacal apertures are large 
and rounded and are occasionally but not 
always found on the free ends of the lobes. 
The test is firm and gelatinous, without sand. 
There is a central gelatinous core of test 
surrounded by specially extensive common 
cloacal spaces. The zooids are small and num- 
erous, closely packed in the outer layer of test. 
There are 8i coils of the vas deferens around 
a single undivided testis follicle. 
Remarks: This species appears to be limited 
to the more temperate waters of Australia 
and New Zealand extending north only to 
N .S. W. on the east coast of Australia. The 
spicules are usually present in the surface test 
at the level of the zooids but are often absent 
in the remainder of the test. In one of the 
present colonies spicules are absent entirely. 
The relationship of the present species to Z>. 
spongioides is close. Both have a firm gel- 
atinous test and a similar common cloacal 
system. In both the spicules are usually absent 
from the central test core and form a layer 
only at the level of the branchial siphons. In 
both they are stellate with about seven conical 
pointed rays. The species appear to be distin- 
guished only by the absence of a superficial 
bladder cell layer in D. lambitum and by the 
larvae which, in the latter species, has sub- 
divided lateral ampullae. Generally the col- 
onies of D. lambitum are higher than those 
of D. spongioides. D. spongioides has been 
recorded from the tropics but D. lambitum 
has not. Tt is possible that both species repre- 
sent different stages in development of a single 
species, however, additional specimens, to- 



gether with larvae will be needed to resolve the 
question. 

Didemnum skeati (Sollas) 

Hypurgon skeati Sollas, 1903, p. 729. 

Didemnum psammatodes var. skeati; Michaelsen, 1920, 

pp. 22, 27 and synonymy. Hastings, 1931, p. 95, 

Kott, 1962, p. 326. 

New Records: Western Port (Crawfish Rock; 
Eagle Rock). 

Distribution: Malaysia; Indian Ocean; Vict.: 
Flinders; Qd.: Moreton Bay, Sarina, Low Isles; 
Torres Strait: Possession Island. The species 
has not been recorded from Western Australia 
but in view of its Indian Ocean occurrence 
could be expected to occur there. 
Description: A large number of extensive 
sheets, blackish in colour owing to embedded 
balls of mud throughout the test. Small groups 
of spicules, as previously described, are pre- 
sent over each branchial aperture. The cloacal 
canals are thoracic. Zooids are very small. 
Remarks: The specimens conform completely 
with previous descriptions. In view of the 
constant nature of this form and its consistent 
differences from D. psammatodes, it has been 
elevated to specific rank. 

Lissoclinum fragile (Van Name) 
(Fig. 26) 

Diplosomoides fragile Van Name. 1902, p. 570. 

Lissoclinum fragile', Eldredge, 1967, p. 245 and syno- 
nymy. 

? Diplosoma (sic) caulleryi Ritter and Forsyth, 1917, 
p. 489. 

? Lissoclinum caulleyri; Van Name, 1945, p. 114. 

? Lissoclinum marpum Millar, 1953. p. 301. 

? Lissoclinum bilobatum Millar. 1955, p. 180. 

? Lissoclinum japonicum Tokioka, 1958, p. 73. 

? Lissoclinum notti Brewin, 1958, p. 457. 

Neyv Records: Western Port (Eagle Rock) 
Distribution: West Indies; ? East Africa; ? 
South Africa; ? Japan; Pacific; ? California; 
? New Zealand. Except for Lissoclinum fragile 
Van Name and Lissoclinum caulleryi Ritter 
and Forsyth, the suggested synonyms are known 
only from single records. The present speci- 
men is the only record from Australia. The 
lack of records may be explained bv the brittle 
nature of the very thin investing colony which 
is removed from the substrate only with the 
greatest difficulty. The species is probably 



ASCIDIAN FAUNA OF WESTERN PORT 



71 



circum-tropical and extends into temperate 
regions of both the northern and southern 
hemispheres. 

Description: The colony is thin and extensive, 
investing a very large specimen of Ascidia 
sydneysis. There are pinkish brown pigment 
cells in the surface test. Spicules are very 
dense throughout, and the colony is very 
brittle. Spicules do not line the branchial lobes 
and are absent from a circular area in the 
region of the branchial aperture through which 
the interior of the colony is visible. They are 
small, 02 to 03 mm in diameter, burr-like 
with many flat ended rays. The cloacal cavity 
is mainly thoracic, primary canals sometimes 
extend to the abdominal level but never pos- 
terior to the zooids. The abdominal portion 
of the zooids is embedded in the basal test 
which is very solid and hard owing to the 
density of spicules. The thoraces cross the 
cloacal cavity in independent test sheaths that 
are interrupted over the dorsal surface and 
most of each side of the thoraces they en- 
velope, A very large 'flap like' lateral organ 
is present supported on the edge of the test 
sheath near the ventral border of the zooid 
and overlapping the branchial sac opposite the 
interval between the third and fourth rows of 
stigmata. There are four rows of stigmata with 
eight stigmata in each row. There are two 
testis follicles with a straight vas deferens, 
hooked proximally around between the two 
testis follicles. 

Remarks: The synonymy suggested above was 
first indicated by Kott (1962). The specimens 
all have a deeply indented atrial aperture, a 
similar two dimensional cloacal system, and 
similar spicules within the same size range 
although there is some variation in their den- 
sity and arrangement. The lateral organ is 
usually present and is flap-like and opposite 
the third to fourth rows of stigmata in all cases 
except in L. fragile; Tokioka, where it appears 
to be elliptical, not supported on the edge of 
the test sheath and opposite the second row 
of stigmata. In L. fragile; Eldredge, it is des- 
cribed as a "small flap-like" organ and in 
L. marpum Millar it is also small. In L. fragile 
Van Name it is not always present and hns 



not been described at all for L. notti Brewin. 

Its presence and degree of development is, 
therefore, apparently variable and its use as 
a distinguishing character would not in any 
case resolve the taxonomy of the forms in- 
dicated above on any rational geographic 
ground. 

Larvae have been described for L. notti 
Brewin and L. fragile; Eldredge. They are 
identical in size and form although some of 
Eldredge's specimens had a layer of small 
opaque particles surrounding the larval body 
similar to the particles described for L. os- 
irearium; Kott (1962). 

The present species differs from L. ostrear- 
ium Michaelsen and L. molle Herdman in the 
absence of a three dimensional cloacal system. 

Lissoclimim osfrearium Michaelsen 

Lissoclinum osfrearium Michaelsen, 1930, p. 526. 
Kott, 1962, p. 308 and synonymy. 

New Records; Western Port (Crawfish Rock; 
Flinders Jetty). 

Distribution: W. Aust.: Rottnest Island; S. 
Aust.: St. Vincent Gulf; Qd.: Great Barrier 
Reef. 

Description: The colonies are very thin invest- 
ing and rather delicate. There are some black 
pigment particles in some parts of the colony. 
There is a thin layer of surface test. The basal 
test is thicker, sometimes enclosing abdomina 
but more often clumps of zooids are anchored 
to the basal test by a single narrow strand of 
test. The common test then subdivides to 
enclose each zooid in an independent test 
sheath for almost its whole extent and anchor- 
ing it anteriorly to the surface test. Each in- 
dividual test sheath is interrupted dorsally to 
expose a large part of the dorsal surface and 
sides of the branchial sac to the extensive 
common cloacal cavity. The spicules are dis- 
tributed in varying density in different parts 
of the colony and are often almost entirely 
absent. They are less dense in the surface 
layer than in the remainder of the test. The 
spicules are 0025 to 003 mm in diameter 
and are characteristic, with a large number 
of flat-ended rays. There is a small, flap-like 
lateral organ opposite the interval between 



I \ I'A'I'KICIA KOTT 

the third and fourth rows of stigmata. No developed and zooida tend to remain in 

embryos are present in thc&e Victorian colonies, clumps. This also gives the colony a firmer 

Remarks: The atrial aperture and the lateral consistency. 

organ are similar to ihose of L. fragile. They 

arc distinguished only by the colony which ,^. , . *„ , A • i 

' ■ , r , Dinlosoiiiii niyncn MaeDonald 

demonstrate!! a maximum development oi the 

cloacal system as in other species of Ltssoc- Diplosoma rayneri MaeDonald, IKV>, p. $73. 

/ i / »■ / Ltptocllnurn i Leptocllnuni ) rayneri: Koii, 1966, p. 

Imum and Uiplosoma. \ tH) v * ' 

Eldrcdgc ( 1967) drew attention ^o the fact Diplosoma Hstgranum; Rowd, 1966, p. 458 and syno- 
nymy. 
Diplosoma macdonaldt\ Eldredge, 1967, p. 231. 



that the difTcrence in the shape of the stigmata 

is probably nol a valid character to distinguish 

/ fragile and L, ostreartum, it is possible New Records; Western Port (Crawfish Rock). 

that the shape of the stigmata is affected by Distribution: Cosmopolitan (see Rowc, L966). 

the extent to which colonies with dense Spi- Description: Typical delicate colonics. Veg- 

cnles retain their original shape when pre etativc reproduction in progress. No mature 

served in formalin so thai the branchial site gonads observed. 

is maintained in an extended condition by the 

rigid test „ . . ■ • ■ i tl 

Polysyncraron ormculum Kott 

Polysyncraton orblculum Kott, 1962. p. 301. Kott, 

i)i|>ioM>m;i transluctdum (Hartmcyer) 1872a, p. 21, 

(Figs, 27, 28) New Records: Western 1*011 (Crawfish Rock). 

Uptocllnum transiucidum Hartmeyer, 1909. p. 1490 Distribution: W. Aust.: Rotlnesl Island; S. 

Diplosoma transiucidum', Kott, 1962, p. 306 and Ausl.: St. Vincent (mil. 

synonymy, Description: The colonies are small and in- 

New Records: Western Port (Eagle Rock). vesting with the usual circle o\' large vesicular 

Distribution: W. Aust.: Oyster Harbour, Al- eells around each branchial aperture. There is 

bany; N. W. Aust.; Indonesia. B Single layer ot spicules in the surface test. 

Description: The colony is lour,, narrow and interrupted by these large vesicular cells. The 

irrcgulai investing a worm tube. It has a zooids, with the red brown pigment In them, 

transparent test that is fairly soft but tough give to the colony a pinkish brown colour. 

and not jelly like. The basal test is extended Sonic-times the vesicular cells are so large 

upwards in a lamella along, the mid longUud- that they are almosl eonlluent. Because these 

iual axis Of the colony. Strands of test from cells are transparent and interrupt the distii- 

both sides o\' Ihis lamella support clumps o[ bulion o[' Spicules the surface of the test 

ZOoids, The common test then subdivides to appears to be pitted, or, when they arc almost 

form tesi sheaths supporting the thorax of confluent, it appears lo be depressed into a 

each zooid independently at the surface. The narrow trough or furrow around each opening 

zooids are fairly large with the thorax about 2 so that the apertures are at the apices o\' 

nun long,. There are four rows o\~ about It) apparent mounds over each zooid. The spicules 

Stigmata. A large pari oi the branchial sac ftW 02 to 003 mm in diameter and are 

is exposed through the wide atrial opening, regularly stellate with about eight conical rays 

There is a long rectum. Oesophageal buds in optical section. 

are present, but gonads were not distinguished. The cloacal canal is shallow and thoracic, 

Remarks: The species is distinguished from D. the zooids small and completely embedded. 

rayneri by its firmer lest and, although the Gonads were not detected. 

cloacal system is typical o\' the genus with Remarks: The form o( the colony and arrange 

long test strands anchoring the /ooids basallw meat of spicules has been used to determine 

the secondary cloacal spaces are not so well this species. 



ASC1DIAN FAUNA OF WESTERN PORT 



73 



Polsyncraton victoriensis n. sp. 

(Fig. 29) 

Type location: Western Port (Crawfish Rock, 
8 m, on Ecklonia holdfasts) Holotype, Nat- 
ional Museum of Victoria No. H. 171. 
Description: The colony forms a thin invest- 
ment over weed. It is a rather dirty whitish 
colour in formalin. There is a layer of bladder 
cells superficially over the top of each zooid. 
Between the zooids, however, spicules invade 
the superficial layer of test which stands out 
as spicule filled ridges between the zooids and 
gives an irregular and rather angular appear- 
ance to the colony. Zooids thus appear to 
open into the base of furrows on the surface. 
There is a very shallow common cloacal 
cavity. Spicules are stellate, 003 to 006 mm 
in diameter with only five conical rays in 
section. They are arranged evenly throughout 
the test. Zooids are small. There are four rows 
each of six stigmata in the branchial sac. There 
are 4] coils of the vas deferens around three 
to four testis follicles. 

Remarks: The arrangement of spicules is the 
same as that described by Hastings, 1931 for 
P. magnetae. Hastings species, however, has 
smaller spicules, fewer turns of the vas defer- 
ens, more testis follicles and more stigmata. 

Plialhisia dcprcssiuscula (Heller) 

Ascidia dcprcssiuscula Heller, 1878, p. 5. 
Piiallusia; dcprcssiuscula; Kott, 1972a, p. 23 and 
synonymy. 

New Records: Western Port (Flinders Jetty; 
Tankerton Jetty); Port Phillip Bay (Hobson's 
Bay; artificial reef). Portland Harbour 6-12 
metres, on rocks forming jetty. 
Distribution: Ceylon; Indonesia; Arafura Sea, 
Philippines; circum-Australia. 
Description: The present specimens fall within 
the range previously indicated for this species. 
Large specimens from Portland Harbour are 
black; while the smaller specimens and speci- 
from Flinders jetty, up to 20 cm in length and 
fixed by the whole of the left side, are brown- 
ish and translucent. One of these large speci- 
mens is completely invested with Lissoclinitm 
fragile. 



Ascidia sydneyensis Stimpson 

(Fig. 30) 

Ascidia sydneyensis Stimpson, 1855 (? pari), p. 387. 
Kott, 1972a, p. 24 and synonymy; 1972, p. 182: 
1972c, p. 237; 1972e, p. 49. 

New Records: Western Port (Tankerton Jetty, 
Flinders Jetty, Crawfish Rock, Eagle Rock); 
Port Phillip Bay (Williamstown, Hobson's 
Bay; artificial reef). 

Distribution: West Jndies; South and east 
Africa; Indian Ocean; Indonesia; circum-Aus- 
tralian. The species is apparently circum-polar 
in tropical and temperate waters of the south- 
ern hemisphere although it extends north of 
the tropics only to Japan. 
Description: Large specimens at least 10 cm 
long and up to 20 cm are available from all 
stations, fixed by the whole of the left side. 
The apertures are on the usual short cylindri- 
cal siphons, the branchial aperture is always 
turned to the left toward the substrate. The 
atrial aperture from half way down the body 
is turned to the right or directed anteriorly 
along the dorsal surface. Sometimes the whole 
antero-dorsal part of the body is turned over 
to the left. The test of these large specimens 
is firm and gelatinous and slightly leathery 
superficially except along the left side where 
it is fixed to the substrate and there it is very 
thin. Two of the four large specimens from 
Flinders jetty are completely covered by in- 
vesting didemnids. On one specimen there is 
a colony of Didemnum posteriorly and a col- 
ony of Tridemnum cyclops anteriorly, which 
leaves only the branchial aperture free. These 
colonies overlap one another across the upper 
surface in a line with the atrial aperture which 
is also left free. On the other specimen the 
upper surface is completely invested with a 
colony of Tridemnum cyclops. There the 
branchial aperture is free but the atrial aperture 
is covered and the excurrent stream from the 
Ascidia is apparently directed along a groove 
in its test to the left of the line between the 
atrial and branchial siphons and underneath 
the encrusting didemnid. The gut, in all these 
specimens, is filled with mud and the branchial 
sac is occluded by the distended gut. 



74 



PATRICIA KOTT 



Ascidia gemmata Sluiter 

Ascidia gemmata Sluiter, 1895 r p. 177. Kott, 1972a, 

p. 26 and synonymy. 
Ascidia thompsoni; Kott, 1975, p. 10. 

New Records: Port Phillip Bay (Mornington 
Pier). 

Distribution: Pacific; Malaysia; Indonesia; 
Arafura Sea; circum- Australia. The species 
thus appears to have a wide range in the Indo- 
Pacific area. 

Description: Two specimens are available, 
about 20 cm long. The branchial aperture is 
terminal on a short siphon. The atrial aperture 
is sessile, two-thirds of the way along the 
dorsal surface. There is a long furrow extend- 
ing along the right side of the body and the 
atrial aperture is directed into that furrow. 
The animal is fixed by the whole of the left 
side. The test is firm, gelatinous and smooth 
on the surface. 

Internally the siphons are more conspicuous. 
There are 60 branchial tentacles, the dorsal 
tubercle fills the peritubercular area. The pre- 
branchial region is minutely papillated. The 
dorsal lamina has the usual strong ribs on 
both sides, each rib terminating in a minute 
pointed tongue to form a marginal fringe. The 
dorsal gland and ganglion are one third of the 
distance down the body. There are intermediate 
papillae in some parts of the branchial sac 
in addition to the papillae at the junction of 
the transverse and parastigmatic vessels. The 
oesophageal opening is just posterior to the 
base of the atrial opening and branchial sac 
extends posteriorly to it. 

Remarks: The species has not yet been record- 
ed from Western Port Bay but will, very 
likely, be found to occur there. 

BotrvIIoides leachi (Savigny) 

Botryllus leachii Savigny, 1810, p. 7. 
Botrxlloides leachi; Kott, 1972a, p. 29 and svnonvmv; 
1972b, p. 185; 1972d. p. 253. 

.Yen- Records: Western Port (Crawfish Rock); 

Portland Harbour (6 metres, on rocks from- 

ing jetty). 

Distribution: The species is recorded from the 

north Atlantic the Mediterranean and Red 

Sea, South Africa, and Indo-Australia to New 

Zealand. The species does not extend into the 



south Atlantic. In view of its south African 
occurrence could be expected to occur more 
widely in the Indian Ocean than its present 
records suggest. 
Description: As previously described. 

BotrvIIoides nigrum Herdman 

Botrxlloides nigrum Herdman. 1886, p. 50. I 
1972c, p. 238 and synonymy; 1972d, p. 252. 

New Records: Western Port (Crawfish Rock; 
Eagle Rock). Port Phillip Bay (artificial reef; 
Mordialloc). Cape Nelson (near Portland; 5 
metres). 

Distribution: West Indies; Red Sea; Ceylon 
(?); South and East Africa; S. W. Aust.; 
Vict.; N. S. \V.; Qld. The species is not as 
commonly recorded as other Botrylloides spp. 
Remarks: The double row systems, widely 
spaced in irregular lamellae are distinctive and 
it seems unlikely that the species could be 
misidentified. Distribution is not cireum-polar, 
as it has not been recorded from the West 
African coast, nor from the Pacific Ocean. 

It may be that there are two species repre- 
sented, one from the Atlantic and one from 
Indo-Australian waters. 

Symplegma viride Herdman 

Symplegma viride Herdman, 1S86. p. 144. Michaelsen, 
1918, p. 101 and synonymy. Kott, 1952. r. 252 and 

further synonymy; 1964. p. 129. 

New Records: Western Port (Crawfish Rock. 
growing on basal surface of fronds of Didem- 
num patulum, encrusting a lump of tar). 
Distribution: West Indies, Red- Sea. West In- 
dian Ocean, Ceylon. Malaysia, the Phiiiptaes 
and circum- Australian from Shark Bay (\Y. 
Aust.) and south across the southern coast 
to Thursday Island (off N. E. Australia). 
Remarks: Colonies of this species appear to 
compete with didemnids for available space 
and occasionally overgrow the borders of a 
didemnid colony. 

Amphicarpa diptycha (Hartmeyer) 

(Figs. 31-34) 

Distomus diptychos Hanmeyer, l^t^. p. 87. 
Ampicarpa diptycha: 19*"2e. p. 50 and further syno- 
nymy. 
Stolonica australis Michaelsen. 1°2", p. 202, Kott, 

1972a. p. 28 and further svnonvmv; 1972b, p. 1S ; ; 
1972c, p. 252. 



ASCIDIAN FAUNA OF WESTERN PORT 



75 



New Records: Western Port (Crawfish Rock). 
SSW. of Cape Grant (220 to 275 metres, on 
a large stone); South-east of Portland (166 
to 220 metres). 

Distribution: North to south-western Australia; 
S. Aust.; Vict.; Tas. (d'Entrecasteaux Chan- 
nel). The species is known from 12 to 24 
metres depth. 

Description: Upright oval to elongate individ- 
uals are joined to a basal membrane or on a 
short stalk from basal stolons investing sponges 
or stones. In some colonies individuals are 
tightly packed to give a cauliflower-like appear- 
ance. The test of adjacent zooids is not con- 
fluent, however, and the zooids in the colony 
are connected with one another only by the 
basal stolon or membrane and by the adherence 
of adjacent zooids to the same sand particle. 
The test is covered with sand. Posteriorly where 
it joins onto the basal membrane, the test may 
be flattened to form a wide pseudo-stalk. 
Zooids are up to 8 mm tall and 3 mm in 
diameter. The branchial apertures are sessile 
and a short distance down the dorsal surface. 
The musculature of the body wall is strong 
especially on and around the siphons but pos- 
teriorly becomes weaker. The branchial folds 
are high and overlapping. There are 19 rows 
of stigmata. The gut forms the usual short 
loop across the posterior end of the body and 
the rectum continues anteriorly at right angles 
to the loop. The stomach is pyriform, ex- 
panded towards the pyloric end. There are 
18 longitudinal folds in the stomach wall and 
on the lateral aspect these folds terminate 
against the sutureline and appear to be oblique 
rather than longitudinal. The gastric caecum 
is continuous with the suture. It extends to- 
wards the pole of the gut loop and curves 
around just distal to the pyloric end of the 
stomach. It is tightly held against the stomach 
by a body wall membrane. It is of variable 
length sometimes curved or hooked or forming 
one complete spiral. There is the usual liga- 
ment extending between the intestine and 
pyloric end of the stomach enclosing a flat 
topped endocarp in the pole of the gut loop. 
There is another similar endocarp between the 



Fl 



oesophagus and the end of the intestine where 
the rectum curves anteriorly. 

The anus terminates in two rounded lips. 
Bisexual gonads are present consisting of a 
large ovum and a wide short oviduct with a 
small male follicle beneath the ovum and the 
body wall, the vas deferens curving around 
to open on top of the oviduct. These gonads 
extend in a single row along the middle of 
the right side of the body and occasionally 
further towards the ventral margin. On the 
left they usually extend in a line obliquely 
across the anterior border of the gut loop and 
along the ventral border of the body. Ovaries 
only occur together with testis follicles, al- 
though occasionally the male gland appears 
to be spent or not to be mature. Numerous 
testis follicles, however, are often scattered 
in clumps antero-ventrally and postero-dor- 
sally at either end of the right row of the 
bisexual gonads and on the left of the body 
they extend around the ventral margin anterior 
to the stomach and ventral to the bisexual 
gonads. This condition appears to occur in 
fully developed mature zooids. In less well 
developed zooids the unisexual testis follicles 
are not present and the testes are confined to 
the bisexual organs as described. Larvae are 
present in colonies from Crawfish Rock. They 
have the usual photolith and a circle of 16 
ampullae anteriorly from the centre of which 
three simple papillae diverge. The larval test 
has a foamy vesicular appearance. 
Remarks: A large number of zooids have 
been examined in order to resolve the con- 
fusion between this species and Stolonica 
australis caused by the variable length of the 
gastric caecum, variation in development and 
position of the gonads and variation in the 
development of the colony. The essential 
differences between Stolonica australis and 
Amphicarpa diptycha (viz. the density of the 
zooids in the colony and the position of gonads 
on the body wall) are both extremely variable 
and are related to the condition and age of 
the colony and the maturity of the gonads, 
and the zooids. Similarly the extent to which 
the basal stolons associated with S. australis 
have fused to form a basal membrane and the 



76 



PATRICIA KOTT 



stalk of the zooid has expanded to become 
confluent with that of adjacent zooids prob- 
ably depends on the maturity of the colony. 
It is apparent from the zooids in the present 
collection, all from the same location, that 
no essential difference separates these species. 
In the present specimens there is an encap- 
sulated parasitic copepod in the peribranchial 
cavity of most individuals. 

Polyandrocarpa lapidosa (Herdman) 

Goodsiria lapidosa Herdman, 1899, p. 99. 
Polyandrocarpa lapidosa', Kott, 195 2, p. 250; 1972b, 
p. 184; Millar, 1963, p. 730. 

New Records; Port Phillip (Survey area 5; 

Popes Eye, Port Phillip Heads). 

Distribution: Port Phillips Heads, Western Port. 

New South Wales (Port Jackson). 

Description: Colony forms a large flattened 

lobe with zooids opening around both sides. 

The zooids conform to previous descriptions 

of this species. 

Polycarpa thelypanes (Sluiter) 

Srylea thelypanes Sluiter, 1904, p. 68. 
Polycarpa thelypanes; Kott, 1952, p. 238. 

New Records: Western Port (Flinders Jetty). 
South to south-east of Portland Harbour, 166 
to 220 metres. 

Distribution: Philippines; W. Aust.: Albany. 
Kott, (1952) suggests that the species might 
have been introduced to Australia with Jap- 
anese oysters (Ostrea gigas) with which it was 
taken. The present records, however, suggest 
that this sandy inconspicuous species may 
have a wider range than previously recognised. 
Description: One small specimen is available 
from Flinders Jetty growing on the test of a 
specimen of Ascidia sydneyensis. A single 
small specimen and four large specimens are 
available from off Portland Harbour. The test 
is very stiff and impregnated with sand. The 
body wall is not very muscular and closely 
adherent to the inner surface of the thin, brittle 
test. The specimens are either dorso-ventrally 
flattened or laterally flattened and lie on the 
substrate on their right side. They are not, 
apparently, fixed to the substrate but lie freely 
on it. The branchial aperture rises from the 
anterior end of the body and is directed at 



right angles to its long axis. The atrial aper- 
ture rises from about half way along the 
dorsal surface. The test is tough leathery and 
whitish. It is impregnated with sand every- 
where except on the short cylindrical siphons. 
The siphons are longitudinally furrowed and 
entirely free of sand. In laterally flattened 
specimens the branchial aperture is turned 
over to the right towards the substrate. There 
is a simple U-shaped dorsal tubercular 
opening in a large peritubercular area. The 
body wall is closely adherent to the test and 
has very delicate though strong muscles that 
radiate from the siphons but fade out on the 
body. The branchial sac has four very low 
rounded folds on each side of the body with 
wide spaces between them. There are about 
15 internal longitudinal vessels on the folds 
and 9-12 between the folds. There is a very 
short, rounded stomach and the gut forms a 
wide open arc opening by an eight lobed anus 
into the base of the atrial aperture. The gonads 
are intermediate between long styelid-type 
gonads and shorter polycarps. There are about 
eight on both sides of the body more or less 
in a row near the endostyle with others irreg- 
ularly scattered over the body wall. Sometimes 
the gonads are slightly curved and bent. 
Remarks: The form and arrangement of gon- 
ads, the course of the gut and the rigid, thin 
and stiff test characterise this species. It is of 
interest that the siphons remain the only con- 
tractile part of this animal, the remainder of 
the body wall is closely adherent to the test. 
The plane along which the animal is flattened 
also appears to be variable and a consequence 
of its position on the substrate. The flattening 
of the body contributes to the orientation of 
the branchial aperture toward the substrate 
and the atrial aperture away from the sub- 
strate. 



Cnemidocarpa etheridgii (Herdman) 

(Fig. 35) 

Stycla etheridgii Herdman, 1899, p. 38. 
Cnemidocarpa etheridgii; Kott, 1972a., p. 31 and 
synonymy; 1972d, p. 253. 

New Records: Port Phillip Bay (Mornington; 



ASCIDIAN FAUNA OF WESTERN PORT 



77 



Port Phillip Heads). South to south-west of 
Cape Grant, (221 to 275 metres). 280° from 
Cape Nelson, (220 to 294 metres); Portland 
Harbour South to south-east of Portland Har- 
bour (166 to 278 metres). 
Distribution: W. Aust.: Triggs Island (north 
of Fremantle); around the south coast of 
Australia; Tag.; d'Entreeasteaux Channel; 
N. S. W.: Port Jackson, Port Stephens; Qd.: 
Moreton Bay. 

Description: The test is whitish and tough and 
leathery, but thin and paperlike in appearance. 
The branchial aperture is terminal and the 
atrial aperture one third to half way down the 
dorsal surface. Both apertures are sessile. The 
body is more or less conical in outline from 
the wide rounded basal or posterior portion 
and narrowing to the terminal branchial aper- 
ture terminally. Posteriorly, the test may be 
produced into numerous tough root-like pro- 
jections or there may be a single stalk. The test 
around the apertures is usually longitudinally 
furrowed or sometimes, when they are with- 
drawn, is transversely wrinkled. The body 
wall is closely adherent to the test. The dorsal 
tubercle is large and protruberant and almost 
fills the peritubercular area. There is a fairly 
wide, smooth margined dorsal lamina. The 
branchial sac has 4 high, almost overlapping 
folds on each side of the body. There is a 
single large gonad in the centre of the body 
wall on both sides of the body, sometimes 
enclosed in endocarp-like thickening of the 
body wall. The gut forms the usual long, 
narrow, curved loop and also encloses an 
endocarp which may cross from the left to 
the right side of the body behind the branchial 
sac. 

Remarks: This species demonstrates a wide 
range in size up to 11 cm in length. It is 
characteristically large and rounded posteriorly, 
with a typical tough whitish thin test. Intern- 
ally the high branchial folds, the narrow 
curved gut loop and the embedded gonads are 
distinctive. As with Ascidia gemmata this 
species has surprisingly not yet been recorded 
from Western Port, but is likely to occur there. 

F2 



Pyura australis (Quoy and Gaimard) 

australis Quoy and Gaimard 

(Fig. 36) 

Ascidia australis Quoy and Gaimard, 1834, p. 614 
Pyura australis, Kott, 1972b, p. 186 and further 

synonymy. 

New Records: Western Port (Crawfish Rock, 
some fixed to a scallop shell and some with 
the stalk encrusted with Amphicarpa diptycha 
or attached to Ecklonia holdfasts). Port Phillip 
Bay (Portsea). 

Distribution: N. W. Aust.: Geraldton; around 
the southern coast of Australia; Tas.: d'En- 
trecasteaux Channel; S. Aust.: St. Vincent 
Gulf; Vict.: Flinders. 

Description: The present specimens are typical 
although the projections from the test are 
sometimes longer and more pointed than is 
usually the case. The branchial aperture is 
turned downwards toward the substrate and 
protected by large tubercular extensions of the 
test. There are the usual stellate siliceous 
spicules 03 mm in diameter in the test 
and pointed spines, less than 0-1 mm in 
length, lining the siphons. The anal border 
has the usual long, finger-like lobes. 

Pyura cataphracta (Herdman) 

(Figs. 37, 38, 39) 

Cynthia cataphracta Herdman, 1899, p, 31. 

Pyura multiradicata; Kott, 1952, p. 269 (part: not 

Herdman's type specimen of Cynthia multiradicata 

< Pyura spinifcra). 

New Records: Western Port (Crawfish Rock). 
South to south-west of Cape Grant, 220 to 
275 metres. South to south-east of Portland 
Harbour, 160 to 220 metres. 
Distribution: Previously recorded only from 
Port Jackson. 

Description: The species is upright, slightly 
narrowed toward the base. The branchial aper- 
ture and atrial aperture are each on a short 
siphon at opposite extremities of the upper 
surface. Basally the diameter of the individual 
is reduced and root-like extensions from the 
test or a single stalk basally rooted may be 
produced posteriorly. Externally the test is 
grooved and furrowed to varying extents and 
especially anteriorly. The test around the 
apertures is produced into pointed processes 



78 



PATRICIA KOTT 



directed anteriorly around the aperture and 
depending on the extent to which the siphon 
is contracted, sometimes extending across the 
closed aperture. On other parts of the body, 
the test may be produced into small tubercles. 
The surface of the test is granular and very 
hard, impregnated with spherical siliceous 
bodies. These are 0-30 mm in diameter, have 
a granular surface, and are densely packed in 
the superficial test, becoming slightly less 
dense internally. The test is, however, thin 
and very tough throughout. Internally the body 
wall is very thin with very strong longitudinal 
muscle bands radiating from both siphons and 
extending posteriorly to break up into an 
irregular network over the gut-loop and pos- 
terior end of the body. Circular muscle bands 
are thinner but form a more continuous layer 
outside the longitudinal bands on the upper 
half of the body. The test is invaginated into 
the siphonal linings and the same spherical 
siliceous spicules are present in this siphonal 
lining. There are also minute pointed spines 
02 mm long directed toward the aperture. 
The dorsal tubercle is large, a U-shaped open- 
ing directed toward the left with both horns 
turned anteriorly. The branchial tentacles have 
primary, secondary and tertiary branches. The 
prepharyngeal area is fairly narrow. The dorsal 
lamina has short closely-set pointed languets 
arising from the border of a narrow membrane. 
There are six branchial folds on each side of 
the body with 18-21 internal longitudinal 
vessels on each fold and two or three vessels 
between the folds. The gut loop is embedded 
in the body wall and forms a curved open 
loop around the posterior end of the body 
with arborescent liver tubules rising from the 
pyloric region. The gonads on each side of the 
body are broken up into about five pairs of 
polycarp-like sacs either side of a median 
duct. On the left, the gonad is embedded in 
the body wall in the primary gut loop; on the 
right side of the body the gonad occupies a 
corresponding position. The anal border has 
six well developed bifid lobes. 
Remarks: The most outstanding characteristic 
of this species is the very hard "sandpaper- 
like" external test, created by the embedded 



siliceous spicules. The species may also be 
distinguished by the test extensions which 
project especially the branchial aperture. 

The species appears to be most closely 
related to Pyura pachydermatina sub. sp. in- 
termedia Michaelsen (which it resembles in 
the form of the anal border, the size and 
shape of the siphonal spines and in the spheri- 
cal siliceous bodies in the test. It may be 
distinguished by the absence of the typically 
long stalk of P. pachydermatina and by the 
shape of the body with the branchial aperture 
terminal and not adjacent to the stalk as in 
Pyura pachydermatina. The present species may 
also be distinguished by the absence of rod- 
shaped spicules in the test and by its simple 
dorsal tubercular opening which is not con- 
voluted as in P. pachydermatina. The body 
wall is also less muscular and more closely 
adherent to the test, a condition which may 
be associated with the very hard and probably 
more rigid test. 

Pyura scoresbiensis Kott 
Pyura scoresbiensis Kott, 1972a, p. 36; 1972b, p. 187. 

New Records: Western Port (San Remo) 
Distribution: S. Aust: St. Vincent Gulf, Spen- 
cer Gulf, Investigator Strait. 
Description: Specimens comprise an aggregate 
of sandy spherical zooids on long stalks that 
taper basally. Both apertures are close together 
on the upper surface of the individual. The 
body is about 2 cm in diameter. There is an 
investment of sand covering the test externally 
and the bodies of adjacent individuals adhere 
to one another to form the aggregate where 
the surface test contacts and adheres to the 
sand encrusting an adjacent individual. The 
stalks, however, remain free from one another. 
There is a slight ridge between the apertures. 
There are the usual six branchial folds on 
each side of the body. The gut forms a wide 
open loop and the rectum extends anteriorly 
towards the base of the atrial aperture. Arbore- 
scent liver lobes are present in the gastric 
region. The gonads consist of approximately 
11 pairs of polycarp sacs on either side of a 
median duct. They are present in the gut 



ASCIDIAN FAUNA OF WESTERN PORT 



79 






Ascidia sydneyensis 

30— individual showing apertures turned over to- 
ward substrate. 

Amphicarpa diptycha 

31, 32, 33 — variable arrangement of gonads on 
right and left sides of the body. 34 — larva. 

Cnemidocarpa etheridgii 

35 — stalked individual. 

Pyura australis 

36 — spicules and siphonal spines. 

Pyura cataphracta 

37— stalked individual. 38 — sessile individual. 



39 — spicules and siphonal spines. 

Pyura irregularis 

40 — siphonal spines. 

Pyura albanyensis 

41 — siphonal spines. 

Pyura stolonifera 

42 — siphonal spines. 43 — spicules from body wall. 

Microcosmus stolonifera 

44 — siphonal spines. 45 — gut loop and gonads. 

Microcosmus squamiger 

46 — siphonal scales. 



80 



PATRICIA KOTT 



loop on the left side of the body and in a 
corresponding position on the right. 
Remarks: The specimens are smaller than 
those usually reported for this species. There 
is also some variation in body shape which 
is spherical rather than the elongate oval as 
in the typical specimens described from South 
Australian locations. Nevertheless, there is no 
character distinguishing them and it is possible 
that their small size results from a less favour- 
able location possibly at the eastern extent of 
the range of this species. 

Pyura irregularis (Herdman) 
(Fig. 40) 

Cynthia irregularis Herdman, 1882, p. 141. 
Pyura irregularis; Kott, 1972a, p. 38 and synonymy. 
1972b, p. 187. 

New Records: Western Port (Crawfish Rock, 
8 metres EckJonia holdfasts; Eagle Rock; 
Tankerton Jetty); Port Phillip Bay (Morning- 
ton Pier). 

Distribution: S. Aust.; St. Vincent Gulf; Tas.: 
d'Entrecasteaux Channel; Vict.: Port Phillip 
Bay; N. S. W.: Port Jackson. 
Description: The present specimens were col- 
lected in large aggregates together with Micro- 
cosmus squamiger. The long siphons are so 
oriented that they are able to reach the exterior 
despite the other individuals with which they 
are so closely aggregated. 

Pyura albanycnsis Michaelsen and Hartmeyer 

Pyura albanyensis Michaelsen and Hartmeyer, 1928, 

p. 435. 
Pyura jacatrensis; Kott, 1952, p. 273. 
Pyura vittata; Miller. 1960, p. 126; Tokioka, 1952, 

p. 134; 1967, p. 202; Kott, 1964, p. 142; 1966, p. 

300; 1972a. p. 37; 1972c, p. 243. 

New Records: Port Phillip Bay (Mornington) 
Distribution: The species has a wide circum- 
Australian distribution. It has also been re- 
corded from the Palau Is. in the Pacific and 
from Ascension Is. in the southern Atlantic. 
Description: The test is very tough, leathery 
and wrinkled. There are the usual six high 
overlapping branchial folds on both sides of 
the body. The dorsal tubercle is a large, double 
spiral cone completely filling the peritubercular 
area. There are needle-like spines lining the 
siphons about 01 mm in length. There are 



endoearp-like blocks of tissue along the an- 
terior limb of the gut loop. The anal border 
is bilabiate. 

Remarks: This species displays to a very high 
degree the taxonomic problems that are en- 
countered in the Class Ascidiacca. Species 
from many localities and with a wide range 
in characters have been included in the syn- 
onymy and Van Name (1921, 1945), Kott 
(1969, 1972a) and Monniot and Monniot 
(1972, 1974) have successively made attempts 
to clarify the situation. Briefly, specimens from 
the West Indies and the Atlantic, the Indo- 
Pacific and the sub- Antarctic (Macquarie, 
Kerguelen and Marion Islands) with siphonal 
spines variously extending onto the outer wall 
of the siphons and the gonads separated into 
polycarp-like sacs along cither side of a central 
duct have been included in the synonymy. 
Monniot and Monniot (1974) have, most 
recently, separated the subantarctic form as 
Pyura pilosa characterised by the absence of 
endocarps on the gut and gonads. In this 
species the anal border is lobed. The siphonal 
spines are long and one of its most conspicuous 
features. Although not described by Monniot 
and Monniot these spines must be regarded as 
contributing to the diagnosis of the species. 

Indonesian and northern Australian species 
P. jacatrensis Sluiter, 1890; Hartmeyer, 
1919; Michaelsen and Hartmeyer, 1928, were 
included in the synonymy of P. vittata by Kott 
(1969) but were later (1966, 1972a) ex- 
cluded on the grounds that the branchial spines 
(003 mm) were very much shorter than those 
of sub-Antarctic and Australian specimens as- 
signed to this species. The anal border in P. 
jacatrensis is simple or very vaguely lobed 
and there are no endocarps on gut or gonads 
(Monniot and Monniot, 1974). 

The West Indies species share, with speci- 
mens from Australia {P. jacatrensis; Kott, 
1952; P. vittata; Kott, 1972a, c), the Arafura 
Sea (P. vittata; Tokioka, 1952), the Palau 
Islands (P. vittata Tokioka, 1967) and As- 
cension Island (P. vittata; Millar 1960), the 
endocarps along the gut loop that are con- 
sidered to be characteristic of P. vittata by 
Monniot and Monniot (1974). Other speci- 



«3 a' 



33 

o P 

P P= 

*o o 



p 

c 
P_ 



p 

5' 



H 3 * 
CO 0) . 
to J> 

p" P 
~. us 

H 

P 



3^3 J? 

re> f& o - 

O (T> O ►— * 

o 

•-* p — J^ 

00 voS 






3 p .3 

rt a ? 

H-J l-» C*J 

p tfl* 



O ©P 



O 
P 



. O 0) 



p 
o 
p 



CO 

■a 


P era 


P 


a*ft 








q"** 


o 




o 




3 





a*t3 



8 ft 
ft o 

en 3 



O ft 

I s 



P ( 

p •* 

ry 

o' 



WO 

p 5r 

3y»5' 



p 



!-, P 

*g 

P 3 

P HH 
CO 

ft 

P 



\n \n \j-> O u-i . 



\OVO ^O 





^D 




ON <-* --J 


^ 






T^ 


;o 




|_4 


t— > 


> 


a 


vo o 


^o 




On — 


as 




^ 


-J , 


r 1 


■^ 


a 


-. H 

[,.0 


^o 


*-* 


2 

69* 


PjO 0\ 
m f*T* ON 

K P " 


o 


o 



O O K: 

p a o 

B.B.8 
o o 

r-t- rt- 



ft OP 

O • ft 

o 

P © — 

o ■ £r 



o 


en 




3 


oo 


p_ 


ft 




o 


^ > 


P 





g &3 p 
2.2 P 



c/j 



O 



P^a" 
►i rs 



£2. 

o £ 
p •** 



'i s n> 

^ 3 



p p 

p p 



3 g 

CO 

o 

p 



-i y o 

3 3^ 

w. 3 



p p 

P s 



p ^-N p ^< 

P^3 P a* 
G.P a g 

O co a pa 

3<JOa 

1— * p ^, 



er 
o 
P 



"2 
3' 



cro 

fD -1 
>-l CO 
O P^ 



^p 



2 "■ o 

P 113 

P P ti 
D-P ►O 
CO p^ CO 

O 

P 



s > 



?^ 




O fD 




►O E" 




S'^ 




P 3 


P 




P 


CO 


P 




ft 



ft ^ 

h-h P 

p* 

o3 

CD v 






3 






fD 



18 



IHOd N*31S3M dO VMQVd MVIQIDSV 



82 



PATRICIA KOTT 



mens from Australia have been re-examined 
(Pt. Vernon: Kott, 1966; Moreton Bay:Kott, 
1964; Ulladulla, N. S. W.; unpublished and 
Port Gregory, Shark Bay, W. A.: unpublished) 
and in each case these endocarps are found to 
be present. All the Australian specimens 
(Kott 1952, 1964, 1966, 1972a, c) and those 
from the Arafura Sea and Ascension Island, 
have siphonal spines of 01 to 0-275 mm and 
these spines confer an irridesccnt greenish 
tinge to the siphons of preserved material. In 
all cases the anal border is bilabate and plain. 
Specimens from Palau Is. and Japan (Tokioka, 
1949b, 1950, 1953) also have a generally 
plain anal border. However, no endocarps 
have been described. The spicules have rot 
been described and the siphons are coloured 
red. The West Indies species differs from the 
Australian and sub-Antarctic forms in the less 
conspicuous siphonal spines (for which the 
length has not been given: Van Name, 1945 
p. 322 "minute short spines, visible only on 
some magnification"), the anal border is lobed, 
and the siphons are coloured red as in the 
Japanese specimens. C. vittata; Oka and C. 
karasboja Oka (Oka, 1935) are identical with 
the West Indies form in all characters. It is 
apparent, therefore, that there are several 
species involved in this complex each charac- 
terised by a reliably constant assemblage of 
characters and in some cases with an over- 
lapping geographical range, as set out in the 
following Table. 

The specimens set out in the synonymy 
above conform with the description of P. 
albanyensis Michaelsen and Hartmeyer. 

Pyura stolonifcra (Heller) subsp. 
praeputialis Heller 

(Figs. 42-43) 

Cynthia stolonifcra Heller, 1878, p. 10, 
Cynthia praeputialis Heller, 1878. p. 12. 
Pyura stolonifcra f. waia Michaelsen and Hartmeyer, 

1928, p. 433. 
Pyura stolonifcra: Sluiter, 1927, p. 43; Kott, 1952, 

p. 274; MacNae and Kalk, 1958; Kott, 1964, p. 

141; Monniot. 1965, p. 100; Day. 1974, p. 35. 
Pyura praeputialis; Millar, 1963, p. 738; 1966, p. 372. 

New Records; Western Port (Eagle Rock); 

Port Phillip Bay (Mornington; Hobsons Bay; 

Prince George Light). Ram Head (18 miles 



south of Mallacoota, 6 metres). 
Distribution: P. stolonijera stolonijera; Cape 
Province and Natal (South Africa); Mosam- 
bique; Dakar; Morocco. P. stolonijera prae- 
putialis; S. W. Aust.; S. Aust.: Outer Harbour; 
Vict.; N. S. W.; Qd.: Noosa. The range around 
the Australian coast is clearly defined and 
accompanied by a decrease in the size of the 
individuals at either end of this range (south- 
western Australia and Noosa, Queensland). 
Description: A single specimen is available 
from Mornington and a tight aggregate of 
seven individuals from Ram Head. The speci- 
mens are of the usual pillar-like form, maxi- 
mum height 3-10 cm and maximum diameter, 
across the top of each individual, 2-4 cm. In 
the specimen from Mornington the posterior 
end of the test is produced into irregular 
processes for adherence to a rocky substrate. 
The specimens have the usual convoluted 
double spiral slit on the large hemispherical 
dorsal tubercle with the open interval directed 
anteriorly. The dorsal lamina has pointed lan- 
guets, there are six to seven high overlapping 
branchial folds on each side of the body and 
the gut forms the usual curved loop termin- 
ating in an anus bordered by three shallow 
lobes. The liver is large and consists of dense 
arborescent tubules. The body musculature is 
strong and branches of the longitudinal muscles 
from the siphons are inserted into a rim of the 
body wall around the anterior surface. These 
muscles effect the anterior depression around 
the siphons that is characteristic of the species. 
Strong longitudinal muscles extend only half- 
way down the body. The siphonal spines are 
from 0-05 mm to 008 mm long, becoming 
larger toward the aperture. Calcareous spicules 
(as described by Millar, 1962 for South African 
specimens) are present in the ventral part of 
the body wall, in the gut and in the endostyle. 
Gonads are divided into large paired blocks, 
densely arranged along either side of a central 
duct, in the gut loop on the left and in a 
corresponding position on the right In older 
individuals there is a fold of tissue which ex- 
tends across the body in front of the atrial 
aperture effectively cutting off the posterior 
part of the peribranchial cavity as an excurrent 



ASCIDIAN FAUNA OF WESTERN PORT 



83 



chamber. There is also a dense "fur" of pointed 
papillae on the body wall of older specimens 
in which gonads are more or less embedded 
in the body wall. 

Remarks: Kott (1952) has described two en- 
vironmental forms of this species one from 
estuarine localities and one from open coast 
situations. The typical estuarinc form is char- 
acterised by the presence of a sandy invest- 
ment on the test and is usually short; and 
posteriorly the test is produced into a beard 
of sandy roots; while the open coast form is 
pillar-like owing to the posterior thickening of 
the test to form a wide solid gelatinous stand 
of the same or of only slightly lesser diameter 
than the rest of the body, thus raising the in- 
dividual above the substrate. Specimens inter- 
mediate between these two forms are known 
and the present specimen from Mornington 
Pier in which the test is produced into irreg- 
ular processes reflects the variability in growth 
of the test of this species. Tn the open coast 
form the gonads are occasionally broken up 
into a single, rather than double, row of sep- 
arate sacs. Variations in the leneth of the 
branchial tentacles also occur and very small 
branchial tentacles have been observed in 
estuarine forms. These characters are variable, 
however, and cannot be regarded as constant 
differences between individuals from these two 
environments. The differences in the test espe- 
cially appear to result from the individual's 
response to different sets of conditions, viz. 
the solid gelatinous pillar of test associated 
with firm fixation to rocky substrates and to 
adjacent individuals where wave action is 
strong; the development of long rooting pro- 
cesses fixing rounded solitary individuals in 
sandy substrates subjected to less turbulent 
conditions. 

There has been considerable discussion in 
the literature on the relationships between the 
South African P. stolomfera and the Austral- 
ian P. praeputialis. Hartmeyer (1911) sug- 
gested that the presence of a seventh branchial 
fold was exclusive to South African forms. 
There are, however, seven branchial folds on 
each side of the body in the present specimens 



from Ram Head and this character does not 
provide a distinction. 

Nor can siphonal spines be used to distin- 
guish subspecies. Michaelsen and Hartmeyer 
(1928) recognised the subspecies P. stoloni- 
fera waia from Western Australia with siphonal 
spines 009 to 01 mm long, and P. stolonijera 
typica from South Africa and P. stolonijera 
praeputialis from New South Wales (Heller's 
type specimen) with siphonal spines 02 to 
0-024 mm long. The present specimens dem- 
onstrate a wide range in length of siphonal 
spine between 005 and 0-08 mm while the 
spines of typical specimens collected from 
Moreton Bay (Queensland) are 0-03 to 0-04 
mm long. Spicules (branched) in the branchial 
sac of South Africa specimens occur in the 
present specimens from Victoria. 

Millar (1963) distinguished the South Afri- 
can form from the Australian by the absence 
of a sunken area around the siphons and by 
the posteriorly directed dorsal tubercle open- 
ing. Some South African specimens (Millar, 
1955) were also distinct in the presence of 
four test projections around the apertures. 
These differences, however, are not constantly 
expressed (Millar, 1966). In fact Cynthia 
valdiviae Michaelsen and C. herdmani Von 
Drasche (see Michaelsen, 1904), both from 
South Africa are identical with the Australian 
form in these characters and in others. Day 
(1974) has described the depressed area 
around the siphons in South African speci- 
mens. In Australian specimens, the collar of 
test around the siphons is invariably present 
when specimens are contracted (i.e. when pre- 
served or exposed intertidally). It is probable 
that it is not present if individuals are not 
contracted. It was not observed by the present 
author in large in situ subtidal populations at 
Noosa (Queensland) in which the siphons of 
all individuals were fully extended in the feed- 
ing position. Day (1974) p. 36, has also ob- 
served variations in form for the South African 
populations that are identical with variations 
in the Australian forms, viz: "solitary speci- 
mens below tide mark are usually hemispheri- 
cal .. . Specimens from S. African estuaries 



84 



PATRICIA KOTT 



which have root like extensions of the test to 
anchor them in the sand". 

It is most probable, therefore, that in view 
of the general form of the body (and its 
capacity to respond to a wide range of en- 
vironments) and the similarities between such 
features as the dorsal tubercle, the siphonal 
spines, and the occasional presence of branched 
spicules, that two separate species cannot be 
maintained. 

Nevertheless, no test projections from the 
siphonal region have ever been observed in 
the many Australian specimens that have been 
examined; and the relationships are most 
reliably indicated by regarding the Australian 
and South African forms as separate geograp- 
hic sub-species, morphologically distinguished 
only by a degree of variability of certain 
characters in the South African populations, 
which are not variable in Australian speci- 
mens, i.e. the absence of projections from the 
siphonal region (sometimes present in S. 
African specimens). 

Pyura lcpidoderma Tokioka 

Pyura lcpidoderma Tokioka, 1949, p. 10. Kott, 1966, 
p. 299. 

New Records; Port Phillip Bay. 
Distribution: Qd.: Hervey Bay. Japan. 
Description: A single flattened specimen, basal 
diameter 1 cm, heighth 2 mm is available, fixed 
to a Mytilus shell. The siphons are on the 
upper surface and are almost sessile. The sur- 
face layer of test is marked off into polygonal 
scale-like thickenings that are most conspic- 
uous around the apertures where they em- 
phasise furrows along the short siphons. The 
body musculature consists of longitudinal bands 
from both apertures which cross one another 
on the body. There arc six branchial folds 
on each side of the body with closely set in- 
ternal longitudinal vessels. The liver branches 
off the gut in the pyloric region and divides 
into arborescent branches along its length. 
The gut loop is narrow and the anal border 
has shallow rounded lobes, the rectum turns 
anteriorly and is very short. The gonads, in 
the gut loop on the left and in a corresponding 
position on the right, consist of polycarp-like 



sacs along either side of common ducts 
opening at the base of the atrial aperture with 
the anus. 

Remarks: Attention has already been drawn 
(Kott 1966, 1969) to the similarity between 
this species and the Antarctic P. squamata 
Hartmeyer (see Kott 1969, p. 136). The oc- 
currence of the species in Port Phillip does 
to some extent satisfy the discontinuity in the 
records of the antarctic species and the north- 
eastern Australian Japanese species respec- 
tively. It is possible that further collecting will 
demonstrate that their ranges are continuous 
and that only a single species is represented. 

Halocynthia hispida (Herdman) 

Cynthia hispida Herdman, 1881, p. 61. 
Halocynthia hispida; Kott, 1968, p. 77 and synonvmy; 
1972b, p. 189. 

New Records: Western Port (Crawfish Rock; 
some on Ecklonia holdfasts; Eagle Rock). 
Distribution: S. Aust.: St. Vincent Gulf; Tas.: 
Bass Strait, d'Entrecasteaux Channel, Maria 
Island; N. S. W.: Port Jackson. Ceylon; and 
off the west coast of North America and 
Japan, (see Kott, 1968). 
Description: The present specimens demon- 
strate the range from the individuals covered 
with branched test tubercles to those in which 
there are no tuberculous extensions of the 
test, (see Kott, 1968). 

Herdmania inomus (Savigny) 

Cynthia mom us Savigny, 1816, p. 143. 
Herdmania momus; Kott, 1972b, p. 189; 1972c, p. 
255 and synonymy. 

New Records: Western Port (Flinders Jetty, 
Eagle Rock). 

Distribution: N. W. Aust. (Broome); S. W. 
Aust.; S. Aust.; Vict; N. S. W.; Qd., Arafura 
Sea; Indonesia; Fiji; the Palau Islands; Tahi- 
ti; Japan; the Indian Ocean; the Red Sea; 
South Africa; and the West Indies. Kott (1972) 
has drawn attention to the lack of distinction 
between the Indo-Pacific populations of this 
species and those from the Atlantic, H. momus 
pallida (see Van Name, 1945). It is most 
probable, therefore, that H, momus is a cir- 
cumtropical species extending into temperate 
regions in the southern parts of its range, i.e. 



ASCIDIAN FAUNA OF WESTERN PORT 



85 



around the South African and south Aus- 
tralian coasts. 

Description: The specimen from Flinders jetty 
is 20 cm long and completely invested with a 
colony of Didemnum patulum, leaving only 
the apertures free. 

Microcosmus australis Herdman 

Microcosmus australis Herdman, 1889, p. 23. Millar, 
1963, p. 741; 1966, p. 373. Kott, 1972e, p. 53. 

Microcosmus claudicans sub sp. australis; Michaelsen 
and Hartmeyer, 1928, p. 404 and synonymy. Kott, 
1952, p. 288. 

ICynthia solanoides Herdman, 1899, p. 29. 

IMicrocosmus solanoides; Kott, 1952, p. 289. 

New Records: Western Port (Crawfish Rock). 
Distribution: see Kott 1972e 
Remarks: There seems little to distinguish this 
species from M. solanoides Herdman except 
the siphonal denticles which Kott (1952) des- 
cribed as not curved and much smaller than 
those of the present species. Herdman's type 
specimen from Port Jackson is the only record 
of M. solanoides and it is likely that it cannot 
be separated from M. australis. 

Microcosmus nichollsi Kott 

Microcosmus nichollsi Kott, 1952, p. 290; 1972c, 
p. 245 and synonymy. 

New Records: Western Port (Crawfish Rock, 
8 metres, on Ecklonia holdfasts). 
Distribution: S. Aust.: St. Vincent Gulf; Vict: 
Flinders. 

Description: Only a single specimen is avail- 
able, 1 cm long. The surface is sandy and 
both apertures are anterior and almost sessile. 
The usual pockets or valves are present at 
the base of the atrial siphon, and the usual 
spines and scales are present in the siphonal 
lining. There are seven branchial folds on each 
side of the body with one or two internal 
longitudinal vessels between the folds. The 
gonads are separated into three blocks and on 
the left the most proximal section of the gonad 
crosses into the pole of the gut loop. 

Microcosmus helleri Herdman 

Microcosmus helleri Herdman, 1881, p. 54 Van Name, 
1945, p. 349 and synonymy; Kott, 1972e, p. 54 
and synonymy. 
New Records: Western Port (Crawfish Rock, 
among Ecklonia holdfasts; Eagle Rock). 



Distribution: W. Aust.: Cape Jaubert to Fre- 
mantle; S. Aust.: St. Vincent Gulf; Qd.: Great 
Barrier Reef, Gulf of Carpentaria, Torres 
Strait; Malaysia. Portugese East Africa 
(Michaelsen 1918). West Indies (Van Name 
1945). 

Description: The individuals are upright and 
more or less egg-shaped with a terminal 
branchial aperture and the atrial aperture about 
half way down the dorsal surface. Around 
each opening the test is produced into lobes 
which fold over when the aperture is con- 
tracted. Superficially the test is covered with 
long branched hairs that are obscured by a 
coating of sand which they enmesh. Beneath 
this sandy coating the test is very thin and 
brittle. There are four, very strong, tongue-like 
projections into the cavity of the branchial 
siphon at its base. There are three pockets 
at the base of the branchial siphon, formed by 
folds of the siphonal lining and these undoub- 
tedly act as cuspid valves. There are six 
branchial folds on each side of the body. The 
dorsal tubercle is U-shaped with both horns 
turned in and completely fills a fairly shallow 
peritubercular area. The gut forms the usual 
narrow loop and in the pyloric region there 
are dense parallel glandular folds or lamellae, 
joined together by an external membrane, re- 
presenting the liver. The long gonad on each 
side of the body is divided into three separate 
sections, and on the left crosses the intestine 
into the pole of the gut loop. 
Remarks: The hard cartilage-like projections 
at the base of the siphon together with relat- 
ively small number of wide overlapping branch- 
ial folds characterise this species. The test 
lobes around the apertures are sometimes 
simple, but often are well developed, tuber- 
culous or branched. In some specimens there 
is no coating of sand but externally the test 
is very hard and produced into pointed and 
sometimes branched papillae. 

Microcosmus stolonifera Kott 

(Figs. 44, 45) 

Microcosmus stolonifera Kott, 1952, p. 291; 1972c, 

p. 245 and synonymy. 
New Records: Western Port (Crawfish Rock). 



86 



PATRICIA KOTT 



Distribution: Previously recorded from QUI. 
(Moreton Bay) S. Aust. (St. Vincent Gulf) 
and Tas. (Tiny Is.). 

Description: Posteriorly the test of these in- 
dividuals is produced into an irregular root- 
like structure, sometimes long branched and 
sturdy, sometimes there is a double projection. 
Occasionally individuals are aggregated to- 
gether. Both apertures are depressed into the 
upper surface and surrounded by a raised, 
rounded fold of test. The test is very hard, 
thin and stiff with a dense layer of embedded 
sand. There are overlapping curved spines 
lining the branchial siphon 06 mm long on 
their concave side but extending from a long 
base of about the same length. Short, irregular 
languets are sometimes present on the pre- 
pharyngeal band. The branchial tentacles have 
primary, secondary and minute tertiary 
branches. The dorsal tubercle is relatively 
small in the centre of the peritubercular area 
and has a simple, U-shapcd opening, some- 
times with a single horn turned in. There is 
a long, smooth-edged dorsal lamina. There 
are seven high and overlapping branchial folds 
on each side of the body, with 15-20 internal 
longitudinal vessels per fold and six to eight 
stigmata per mesh. The gut forms a narrow 
loop with liver lamellae in the pyloric region. 
Single rounded or sometimes irregular gonads 
are present on each side of the body. The 
gonad on the left is in the secondary gut loop 
and does not extend into the primary gut 
loop. 

Remarks: The rounded fold of test around the 
upper surface enclosing the apertures is not 
always present and depends to some extent 
on the size of the individual. The species is 
distinctive, however, in the presence of the 
gonad outside the primary gut loop and in 
the size and form of the siphonal spines to- 
gether with the high overlapping folds in the 
branchial sac. 

Microcosmus squamiger Michaelsen 
(Fig. 46) 

Microcosmus claudicans sub. s. squamiger Michaelsen, 
1928, p. 405. 

Microcosmus squamiger; Kott, 1972a, p. 43 and syno- 
nymy. 



New Records: Western Port (Crawfish Rock; 
Eagle Rock; Rutherford Channel). Port Phil- 
lip Bay (Mornington Pier, 9 to 35 metres, 
dense rock, vertical and horizontal clumps 
fixed to oyster shells; Williamstown, 5 metres, 
common on rocks). 

Distribution: W. Aust: Shark Bay to Albany; 
S. Aust: St. Vincent Gulf; N.S.W.: Port Jack- 
son; Qd.: Bowen and Rockhamton. Red Sea. 
Undoubetdly there has been great confusion 
between this and related species (see Michael- 
sen and Hartmeyer, 1928) and it is probable 
that there is a wider distribution in the Indian 
Ocean, 

Description: Rounded individuals are present 
in sometimes very large aggregates. The aper- 
tures are about one third of the body length 
apart and sessile. The animals are whitish 
to pinkish-brown, sometimes smooth, occasion- 
ally with some embedded sand. The surface 
is often very uneven and wrinkled, especially 
around the siphons. There are minute over- 
lapping siphonal scales about 001 mm long. 
There are eight to nine overlapping branchial 
folds on each side of the body. The dorsal 
tubercle is always a double spiral cone, the 
gut forms a narrow loop, the gonads are 
separated into three sections and cross into 
the primary gut loop. 

Remarks: This is a common species in those 
areas from which it has been recorded and 
is distinguished by the very small overlapping 
siphonal scales, the dorsal tubercle, the 
leathery test and the absence of a dense, 
sandy coating and the large number of over- 
lapping branchial folds. 

Molgula sabulosa Quoy and Gaimard 

Ascidia sabulosa Quoy and Gaimard, 1834, p. 613. 
Molgula sabulosa; Michaelsen and Hartmeyer, 1928, 

p. 449 and synonymy. Kott, 1972c, p. 248 and 

synonymy. 

New Records: Western Port (Crawfish Rock, 
San Remo; Shoreham). Port Phillip Bay. 
Distribution: Known only from Albany, (W. 
Aust.) and Port Phillip Bay (Vict.) 
Description: Large spherical specimens with 
characteristic hollow test lobes formed around 
the apertures. The individuals often form 



ASCIDIAN FAUNA OF WESTERN PORT 



87 



aggregates. The test is thin but stiff with 
adherent sand. The apertures are close to- 
gether on the upper surface but are not in 
a depressed pit as in M . mollis, and do not 
have a thickened ridge of test extending along 
the dorsal line between them. Each aperture 
is surrounded by a rosette of hollow rounded 
lips of the test, 6 around the branchial aper- 
ture and four around the atrial aperture. 
These fold inwards over the aperture and 
each aperture together with its surrounding 
test projections, occupies a circular depres- 
sion in the upper surface of the body. The 
body wall projects into these hollow lobes. 
The dorsal tubercle occupies the right hand 
side of the peritubercular area and there is 
elongate ganglion slightly to the left. The 
dorsal lamina is rather long. There are seven 
narrow branchial folds on each side of the 
body with only three internal longitudinal 
vessels, one on the edge of the fold and the 
other two ventrally. Stigmata are irregular 
between the folds and between the dorsal 
lamina and the first fold. The infundibula are 
tightly coiled and bifurcate in the summit of 
each fold. The gut forms a long narrow curved 
loop and the plain bordered anal opening 
is at the base of the atrial siphon. There 
are many fine longitudinal liver lamellae at 
the cardiac end of the stomach but at the 
pyloric end there are minute arborescent 
lobes. The kidney is long and curved on the 
right hand side of the body and the long 
tubular ovary extends parallel to its dorsal 
surface. Testes follicles are long narrow and 
deeply lobed at their outer edge tapering to 
the vasa efferentia at the end of the ovarian 
tube where they join into the short vas def- 
erens that extends onto the mesial surface 
of the ovarian tube and opens to the peri- 
branchial cavity there. 

Remarks: This species is distinguished from 
M. mollis by the short vas deferens 
(which in some specimens is turned antero- 
ventrally), by the longer dorsal lamina and 



by the hollow lobes of test that protect the 
apertures. 

Mogula mollis Herdman 

Mogula mollis Herdman, 1899, p. 54; Kott, 1952, 
p. 298; 1964, p. 144; 1972a, p. 45 and synonymy. 
Mogula sabulosa; Kott, 1972a, p. 190, 1972d, p. 

248. 

New Records; Western Port (Crawfish Rock). 
Distribution: S. Aust: St. Vincent Gulf; east- 
ern Australia to Indonesia. 
Description: The specimens are smaller than 
is usual for M. sabulosa and are laterally 
flattened and covered with delicate hairs to 
which sand adheres. There are small test pro- 
jections around the apertures but these are 
not hollow as in M. sabulosa. The apertures 
are close together on the upper surface of 
the body both sunk deeply into the surface, 
with a ridge of slightly thicker test between 
them along the dorsal line. The apertures 
are directed away from one another. The 
body wall has very strong muscles in the 
siphons and around the anterior part of the 
body at the base of the siphons but poster- 
iorly the musculature is weaker. The branch- 
ial folds are broad, overlapping and deeply 
curved and the dorsal lamina is very short. 
There may be as few as 4 internal longitud- 
inal vessels on both sides of each fold. The 
gut forms a long, very deeply curved loop 
which enclosed the gonad on the left side of 
the body. At the cardiac end of the gastric 
region there are short transversely oriented 
liver lamellae but at the pyloric end there 
are longer and longitudinally arranged diver- 
ticulae. The testis follicles are grouped around 
the proximal end of the ovarian tube. They 
are long narrow and wedged shaped in out- 
line, converging to paired ducts which join 
into a long vas deferens that extends along 
the middle of the mesial surface of the 
ovarian tube and opens at the base of the 
oviduct. On the right side of the body the 
testis lobes are accommodated in the curve 
of the kidney and not anterior to it as in 
M. sabulosa. 



88 



PATRICIA KOTT 



I»RW uvtr 



niii»»sTQim_ 




47 — Map of region. 



ASCIDIAN FAUNA OF WESTERN PORT 



89 



CHARACTERISTICS OF THE ASCIDIAN FAUNA 
(Fig. 48) 

Most collecting of the Victorian ascidian 
fauna has been done in either Western Port 
or Port Phillip Bay. It is probable that a 
proportion of the very large number of spec- 
ies recorded from these two locations also 
occur on the open coast where a relatively 
limited range of habitats has been sampled. 

The material reported on by Millar (1966) 
from Port Phillip Bay was collected by bot- 
tom sampler and the species taken represent 
the fauna of the sea floor. Kott (1952, 1957, 
1962, 1963) reported largely on intertidal 
collections; while the present mainly subtidal 
collections have been taken manually from 
rock ledges, caves and from the sea floor 
and represent the larger components of the 
benthic fauna. 

There are 26 species that have been re- 
corded from Port Phillip Bay but do not 
occur in Western Port. Three are probably 
introduced (viz. Ciona intestinalis; Ascid- 
iella aspersa; Kott, 1952; and Styela clava: 
Holmes, 1976). Ritterella assymmetrica 
Millar, 1966 and Protopolyclinum sabulosa 
(Millar, 1963) are endemic to Port Phillip 
Bay. Of the remaining 21 species, 12 are 
within their Australian range; and 7 are tem- 
perate species at the eastern or northern ex- 
tremity of their range (including Polycitorella 
mariae; Millar, 1963, previously recorded 
from South Africa and New Zealand). For 
only three of these species does their occur- 
rence in Port Phillip represent the southern 
limit of their range on the eastern coast of 
Australia. 

There are 45 species that have been re- 
corded from Western Port but have not been 
taken from Port Phillip. Twenty four of 
these are recorded only from Crawfish Rock 
and at no other location and a further 10 
occur only at the adjacent Eagle Rock or at 
both Eagle Rock and Crawfish Rock. Of 
this 34 species one is endemic (Polysyncraton 
victoriensis sp. nov.); Dumus arenijerus pre- 
viously known only from New Zealand is re- 
corded from Australia for the first time; 16 
are in the middle of their range; eight are 



at the southern limit of their range and eight 
are at the eastern limit of their range. Of 
the species that are also recorded from other 
parts of Western Port three are in the middle 
of their range, five are at the eastern end of 
their range and three are at the southern end 
of their range (Fig. 48; Tables 1, 2). 



Comparison of the geographical affinities of 
ascidian fauna of Western Port and Port Phillip 
Bay 



i 


i 

I 








1, 



^ 


Introduced Species 


^ 


Endemic Species 


rum 


Species at southern 
limit of range 


F=l 


Species at eastern 
limit of range 


1 1 


Middle of range 


PP Port Phillip Bay 


WP Western Port 


CR C 


rawfish Rock 




(Western Port) 


ER E 


agle Rock 




(Western Port) 


(WP) Western Port but not 


from CR or ER 


Scale 




CZD 


= 1 species 



PP*WP PP CfUER (WP) 

48 — Histograms showing geographic affinities of 
the Western Port and Port Phillip Bay ascid- 
ian fauna. 

TABLE 1 

Geographical affinities of ascidian species from 
Port Phillip Bay* 



Species 



Geographical 
affinity 



Ciona intestinalis 
Protopolyclinum sabulosa (Millar, 

1963) 
Polycitorella mariae', Millar, 1963 
Distaplia viridis; Kott, 1972a 
Distaplia stylifera; Kott, 1972b 



Introduced 

Eastern limit 
S. Africa; N.Z. 
Eastern limit 
Middle of range 



90 



PATRICIA KOTT 



Middle of range 

Endemic 

Middle of range 
Southern limit 
Eastern limit 
Middle of range 
Middle of range 
Middle of range 
Introduced 
Middle of range 
Eastern limit 

Middle of range 
Southern limit 

Middle of range 
Middle of range 
Middle of range 
Introduced 
Eastern limit 
Northern end of 

range 
Middle of range 
Southern limit 

* A recent reference is given for each species not 
discussed above. 

TABLE 2 



Cystodytes dellechiajei; Kott, 1972b 
Ritterella assymmetrica Millar, 

1966 
Euherdmania australis\ Kott, 1972b 
Aplidiwn solidum; Millar, 1963 
Didemnum lambitum 
Perophora hutchisoni; Millar, 1966 
Ascidia gemmata 
Ascidia aclara; Millar 1963 
Ascidiella aspersa; Millar 1966 
Corella eumyota; Knott, 1972a 
Oculinaria australis; Millar, 1966 
Botrylloides magnicoecus; Millar, 

1966 
Polyandrocarpa lapidosa 
Polycarpa pedimculata; Millar, 

1966 
Cnemidocarpa etheridgii 
Styela plicata; Millar, 1966 
Styela clava; Holmes, 1976 
Astereocarpa cerea; Millar, 1966 
Pyura fissa; Millar, 1966 

Pyura albanyensis 
Pyura lepidoderma 



Geographical affinities of ascidian species from Western Port 11 







M 




c o 










o 

o 


o 


.2 °- 


3 




Species 






o 


o <u 

1—1 crt 




Geographical affinities 












|h 








i-H 

u 




OB 


o 




Podoclavella cylindrica 








X 




Eastern limit 


Oxycorynia pseudobaudinensis n. sp. 


X 








Eastern limit 


Eudistoma pyri forme 




X 








Middle of range 


Polycitor giganteum; Kott, 


1972b 


X 




X 




Middle of range 


Atapozoa mirabilis 








X 




Eastern limit 


Sycozoa cerebriformis 




X 




X 




Middle of range 


Sycozoa pedunculata 








X 


X 


Middle of range 


Pseudodistoma cereum 




X 








Middle of range 


Dumus areniferus 




X 








Single Australian record 


Polyclinum marsupiale 




X 








Middle of range 


Aplidium australiensis Kott 


, 1963 






X 




Eastern limit 


Aplidium parvum Kott, 1963 






X 




Eastern limit 


Aplidium pliciferum; Kott, 


1972a 






X 


X 


Middle of range 


Aplidium depressum 




X 




X 




Southern limit 


Aplidium lobatum 




X 








Southern limit 


Aplidium triggiensis 




X 








Eastern limit 


Aplidium opacum Kott, 1963 






X 




Middle of range 


Synoicium hypurgon 




X 








Middle of range 


Sidneyoides tamaramae 




X 








Southern Limit 


Didemnum moselyi 




X 


X 






Middle of range 


Didemnum patulum 




X 


X 






Middle of range 


Didemnum turritum 




X 


X 






Eastern limit 


Didemnum august i 




X 








Eastern limit 


Didemnum roberti 




X 


X 






Eastern limit 


Didemnum spongioides 




X 








Middle of range 



ASCIDIAN FAUNA OF WESTERN PORT 



91 



Species 



Didemnum skeatii 
Didemnum candidum 
Trididemnum cerebriforme 
Trididemnum cy clops 
Lissoclinum fragile 
Lissoclinum ostrearium 
Diplosoma translucida 
Diplosoma rayneri 
Polysyncraton orbiculum 
Polysyncraton victoriensis n. sp. 
Phallusia depressiuscula 
Ascidia sydneyensis 
Symplegma viride 
Amphicarpa diptycha 
Botrylloides nigrum 
Botrylloides leachii 

Polycarpa thely panes 

Pyura australis 

Pyura cataphracta 

Pyura irregularis 

Pyura scoresbiensis 

Pyura stolonijera praeputialis 

Halocynthia hispida 

Herdmania momus 

Microcosmus australis 

Microcosmus nichollsi 

Microcosmus helleri 

Microcosmus stolonijera 

Microcosmus squamiger 

Molgula mollis 

Molgula sabulosa 



l 

i 




a o 

.2^ 






M 
o 




0< 


Geographical affinities 


£ 

tf 


1 








t- 


S 


a - 


o 




u 


tu 


5.S 


Oh 




X 


X 






Southern limit 


X 








Middle of range 


X 








Middle of range 




X 


X 




Southern limit 




X 






Southern limit 


X 


X 






Southern limit 
Southern limit 


X 








Middle of range 


X 








Eastern limit 


X 








Endemic 






X 


X 


Middle of range 


X 


X 




X 


Middle of range 


X 








Middle of range 


X 






X 


Eastern limit 


X 


X 




X 


Middle of range 


X 




X 




Middle of range 
Southern limit 


X 








Eastern limit 


X 








Southern limit 


X 


X 


X 
X 


X 


Middle of range 
Eastern limit 




X 




X 


Middle of range 


X 


X 
X 






Middle of range 
Middle of range 


X 








Southern limit 


x 








Eastern limit 


X 


X 






Middle of range 


X 








Middle of range 


X 


X 




X 


Middle of range 


X 




X 


X 
X 


Southern limit 
Eastern limit 



* The most recent reference is given for each species not discussed above. 



It is probable that both Port Phillip Bay 
and Western Port provide different habitats 
for ascidian species since only a small group 
of species (the majority of these well within 
the limits of their geographic range) have 
sufficiently unrestricted habitat requirements 
to be present in both locations. Larger num- 
bers of species are recorded from either Port 
Phillip Bay or Western Port but not from 
both. The most striking differences in the 
biogeographic affinities of the ascidian fauna 
at these two locations is the high diversity of 
species and the high percentage of northern 
forms at the southern limits of their range 
that occur in Western Port. These differences 
are not associated with a random distribution 
of habitats since the majority of species have 



been taken at Crawfish Rock and also at 
adjacent Eagle Rock and have not been re- 
corded at more southerly locations in West- 
ern Port. Both these stations are in the north- 
ern section of the bay, where there is comp- 
lete protection from oceanic swell and where 
extensive tidal flats draining into the main 
channel may modify the temperate marine 
environment. This may contribute to the 
diversity of environmental conditions that 
support such a diverse fauna with such a 
high proportion of northern species. 

The most diverse assemblage of species 
has been taken from Crawfish Rock, on a 
30° slope of soft brown coral and occasional 
sandstone boulders. The species are largely 
encrusting aplousobranch forms with vivi- 



92 



PATRICIA KOTT 



parous larvae but there are also small leathery 
oviparous stolidobranch species that produce 
root-like structures and form aggregates. The 
stalked species Pyura australis only rarely 
occurs here, and large stolidobranch and 
phlebobranch forms (Phallusia depressiuscula, 
Herdmania momus, Cnemidocarpa etheridgii, 
etc.) that require smooth and firm surfaces 
for fixation are not present. 

The turbidity of the water and its effect 
on the light intensity at greater depths does 
not appear to limit ascidian distribution at 
this location. 

Similar, though not such dense nor diverse 
associations of species occur at Eagle Rock 
and in the Rutherford Channel. 

It is possible that the less diverse fauna of 
Port Phillip Bay is a result of environmental 
disturbance. The presence there of (probably) 
introduced species Ciona intestinalis, Ascidi- 
ella aspersa and Styela clava (see Holmes, 
1976) may be evidence of this disturbance. 

The occurrence of Dumus areniferus B re- 
win, previously known only from New Zea- 
land, increases the number of temperate spec- 
ies that are known to occur in both southern 
Australian and New Zealand waters (Kott, 
1974). It is unlikely to have been introduced 
on ship's hulls and its Australian occurrence 
may have previously been overlooked. 

STATION LISTS 

WESTERN PORT 

CRAWFISH ROCKS, tidal currents, 5 knots 
? metres: 

Aplidium triggiensis 

Trididemnum cerebriforme 
Intertidal: 

Microcosmus squamiger 

Microcosmus australis 
0-15 metres: 

Sycozoa cerebriformis 

Dumus areniferus 

Aplidum lobatum 

Sidneoides tamaramae 

Didemnum skeati 

Didemnum spongioides 

Amphicarpa diptycha 

Pyura irregularis 

Halocynthia hispida 

Mogula mollis 
8 metres, Ecklonia 'holdfasts': 

Aplidium lobatum 

Didemnum candidum 



Didemnum mosleyi 

Didemnum patulum 

Polysyncraton victoriensis 

Botrylloides leachii 

Pyura australis 

Pyura cataphracta 

Pyura irregularis 

Halocynthia hispida 

Microcosmus helleri 

Microcosmus nichollsi 

Microcosmus stolonifera 

Microcosmus squamiger 
12 to 24 metres: 

Oxycorynia pseudobaudinensis 

Sycozoa cerebriformis 

Eudistoma pyriforme 

Polycitor giganteum 

Pseudodistoma cereum 

Synoicium hypurgon 

Polyclinum marsupiale 

Aplidium lobatum 

A plidium depressum 

Didemnum moseleyi 

Didemnum patulum 

Didemnum turritum 

Didemnum augusti 

Didemnum skeati 

Didemnum spongioides 

Didemnum roberti 

Polysyncraton orbiculum 

Lissoclinum ostrearium 

Diplosoma rayneri 

Ascidia Sydney ensis 

Botrylloides nigrum 

Symplegma viride 

Amphicarpa dytycha 

Pyura australis 

Halocynthia hispida 

Microcosmus helleri 

Microcosmus stolonifera 

Microcosmus squamiger 
13-26 metres: 

Amphicarpa dipytcha 

Pyura australis 

EAGLE ROCK, 15 metres: 

Trididemnum cyclops 
Didemnum mosleyi 
Didemnum patulum 
Didemnum turritim 
Didemnum roberti 
Didemnum skeati 
Lissoclinum fragile 
Diplosoma translucidum 
Ascidia sydneyensis 
Botrylloides nigrum 
Pyura irregularis 
Pyura stolonifera praeptialis 
Halocynthia hispida 
Herdmania momus 
Microcosmus helleri 
Microcosmus squamiger 

RUTHERFORD CHANNEL, fast current, 5 metres: 
Sycozoa penduculata 
Aplidium depressum 
Microcosmus squamiger 



ASCIDTAN FAUNA OF WESTERN PORT 



93 



TANKERTON JETTY, 7 metres: 
Polycitor gigan/eum 
Atapozoa mirabilis 
Sycozoa cerebriformis 
Aplidium pliciferum 
Phalhtsia depressiuscula 
Pyura irregularis 

SHOREHAM: 

Ascidia syneyensis 
Molgula sabulosa 
SAN REMO: 

Pyura scoresbiensis 
Molgula sabulosa 

BALNARRING BEACH: 

Podoclavella cylindrica 
FLINDERS JETTY, 3 metres: 

Oxycorynia pseudobaudine nsis 
Podoclavella cylindrica 
Trididemnum cyclops 
Lissoclinum ostrearium 
Phallusia depressiuscula 
Ascidia sydneyensis 
Polycarpa thelypanes 
Herdmania momus 

PORT PHILLIP BAY 

WILLIAMSTOWN, 5 metres (common on rocks): 
Ascidia sydneyensis 
Mierocosmus squamiger 

HOBSONS BAY, 13 metres: 

Didemnum lambitum 
Ascidia sydneyensis 
Phallusia depressiuscula 

MORNINGTON, 12/10/69, bottom patchy, rock and 
sand, coll. Kevin Duke, 1-2 metres: 
Pyura albanyensis 
Pyura stolonifera praeputialis 
MORNINGTON PIER, 12/10/69, sparse rock, calm 
waters, coll. Kevin Duke: 
Ascidia gemmata 
Cnemidocarpa etheridgu 
MORNINGTON PIER, 12/10/69, dense rock, coll. 
Kevin Duke, 8-11 metres: 
Pyura irregularis 
Mierocosmus squamiger 
YARRA RIVER, 0-3 metres, 28/4/72, No. 3 Oil 
Wharf, Coll. J. E. Watson. 
Ciona intestinalis 
PORTSEA, 15/9/1957: 

Amphicarpa diptycha 
MORDIALLOC BEACH, Nov, 1888, Coll. W.K.; 

Aplidium pliciferum 
Botrylloides nigrum 
ZOOBENTHOS SURVEY, Fisheries and Wildlife De- 
partment, Coll. G. Poore: Off BRIGHTON, 
21/10/69, Station 906, sandy bottom, 10 metres; 
middle of northern part of Gulf, 10/2/69, 
Station 915, silty-clay bottom, 19 metres; of! 
GEELONG, 12/2/70, Station 940, silty clay 
bottom, 8 metres: 



Ascidia sydneyensis 
South of POINT WILSON, 12/2/70, Station 
942, silty clay bottom, 7 metres: 

Ascidia sydneyensis 

Molgula sabulosa 
North-west of PORT ARLINGTON, 11/2/70, 
Station 930, silty clay-sand bottom, 10 metres: 

Ascidia gemmata 
North of PORT ARLINGTON, 18/2/71, Station 
931, silty sand-shell bottom, 15 metres; north- 
west of ROSEBUD, Station 982, sandy bottom, 
18 metres: 

Sycozoa pedunculata 

Cnemidocarpa etheridgii 
East of ST. LEONARDS, 16/2/7 1, Station 960, 
sandy bottom, 10 metres: 

Moluga mollis 

Off BRIGHTON, 9/3/71, Station 1218, sandy 
bottom, 4 metres: 

Pyura lepidoderma 

Pyura stolonifera praeputialis 
Off BRIGHTON, 9/3/71, Station 1226, silty- 
clay bottom, 8 metres; half-way down eastern 
shore of the Bay, 10/3/71, Station 1241, sandy 
bottom; south of SOUTH WERRIBEE, 11/3/71, 
Station 1252, sand-gravel bottom, 4 metres: 

Pyura stolonifera praeputialis 
PORT PHILLIP SURVEY: 

area 5, POPES EYE, 15/5/63: 

Polyandrocarpa lapidosa 
area 30, PRINCE GEORGE LIGHT: 

Pyura Stolonifera praeputialis 
ARTIFICIAL REEF: 

Ciona intestinalis 

Sycozoa pedunculata 

Ascidia syneyensis 

Botrylloides nigrum 

References 

Brrwin, Beryl I, 1952. Ascidians of New Zealand. 
Part VII. Ascidians from Otago coastal waters 
Part II. Trans. R. Soc. N. Z. 79: 452-458. 

, 1958. Ascidians of New Zealand, Part XII. 

Ascidians of the Hauraki Gulf Pt. III. Trans. R. 
Soc. N.Z. 85 (3): 455-458. 

Day, R. W„ 1974. An investigation of Pyura stoloni- 
fera (Tunicata) from the Cape Peninsula. Zoo- 
logica africana 9 (1): 35-58. 

Eldredge, L. G., 1967. A taxonomic review of Indo- 
Pacific didemnid ascidians and descriptions of 
twentv-three central Pacific species. Micronesia 2: 
161-261. 

Gottschaldt, R., 1898. Synascidien von Ternate. 
Abh. Senckenb. Naturf. Ges. 24: 641-666. 

Hartmeyer, R., 1909-11, Ascidien (continuation of 
work by Seeliger). In Bronn, H. G., Klassen and 
Ordnungen des Tier-reiches. Leipzig, 3, suppl.. 
89-98): 1281-1772. (Abstract, repeating lists of 
species by Schepotieff, A., in Arch. Naturgesch,, 
1911, 6: 3-27). 

— , 1911. Die ascidien der deutschen siidpolar 

Expedition 1901-1903. Dt. Siidpolar-Exped. 12: 
408-606. 



94 



PATRICIA KOTT 



, 1913. Tunicata. (In: L. Schultze, Zool. u. 

anthrop. Ergcbnissec Forschungsreise in Sud- 
afrika. Bd 5, Lft 2.) Denkschr, med-natnrw. 
Ges. Jena 17: 125-144. 

, 1919. Ascidien. In Results of Dr. E.Mjo- 

berg's Swedish scientific expeditions to Australia 
1910-13. K. sevnska Vetensk-Akad. 60 (4): 
1-150. 

Hastings. Anna B., 1931. Tunicata. Sclent. Rep. Gt 
Barrier Reef Exped. 4 (3): 69-109. 

Heller, C, 1878. Beitrage zur nahern kenntniss der 
tunicaten. Sber. Akad. Wiss. Wien. 77 (1): 1-28, 
+ P^. 

Herdman, W. A., 1881. Preliminary report on the 
Tunicata of the Challenger expedition. Cyn- 
thiidae. Mogulidae. Proc. R. Soc. Edinb. 11 
(3): 52-88; (4): 233-240. 

, 1882. Report on the Tunicata collected 

during the voyage of H.M.S. Challenger during the 
years 1873-1876. Pt. I, Ascidae simplices. Zool. 
Chali. Exp. 6 (17): 1-296. 

■ , 1886. Report on Tunicata collected during 

voyage of H.M.S. 'Challenger' during years 1873- 
76 Pt. II. Ascidae compositae. Zool. Chad. Exp. 
14 (38): 1-425. 

, 1899. Descriptive catalogue of the Tuni- 
cata in the Australian Museum, Sydney. Cata- 
logue 17: 1-139. 

Holmes, N., 1976. On the ascidian Stvela clava 
in Hobson's Bay. Proc. R. Soc. Vict. 88: 115-116. 

K\ Steven, H. L., 1909. Studies on Tunicata, no. 1. 
Proc, Linn. Soc. N.S.W. 34: 276-295. 

Kott, Patricia. 1952. The ascidians of Australia. 
I. Stolidobranchiata and Phlebobanchiata. Aust. 
J. mar. Freshw. Res. 3 (3): 206-333. 

, 1954. Tunicata. Rep. B.A.N. Z. antarct. 

Res. Exped. I (4): 121-182. 

, 1957. The ascidians of Australia. II. 

Aplousobranchiata Lahille; Clavelinidae Forbes 
and Hanly and Polyclinidae Verrill. Aust. J. 
mar. Freshw. Res. 8 (1): 64-110. 

, 1962. The ascidians of Australia. 111. 

Aplousobranchiata Lahille; Didemnidae Giard. 
Aust. J. mar. Freshw. Res. 13 (3): 265-334. 

, 1963. The ascidians of Australia. IV. 

Aplousobranchiata Lahille; Polyclinidae Verrill 
(ctd.) Aust. J. mar. Freshw. Res. 14 (I): 70- 
118. 

, 1964. Stolidobranch and phlebobranch 

ascidians of the Queensland coast. Pap. Dep. 
Zool. Univ. Qd. 2 (7): 127-152. 

1966. Ascidians of north Australia. Pap. 

Dep. Zool. Univ. Qd. 2 (15): 279-304. 

, 1968. A review of the genus Halocynthia 

Verrill, 1879. Proc. Linn. Soc. N.S.W. 93 (1): 
76-89. 

, 1969. Antarctic Ascidacea. A monographic 

account of the known species based on specimens 
collected under U.S. Government auspices 1947 
to 1963. Antarct. Res. Ser. 13: i-xv, 1-239, figs. 
1-242, pi. 1-3, map, tables— 19. 

, 1971. Antarctica Ascidiacea 11. Collections 

made south of 40° south latitude 1963/67, prin- 
cipally by the U.S.N.S. Eltanin. Antarct. Res. 
Ser., Biology, 16 (4): i-iii, 1-60. 

1972a. The ascidians of South Australia I: 



, 1972b. The ascidians of South Australia II: 

Investigator Strait, and the eastern end of the 
Great Australian Bight. Trans. R. Soc. S.A. 96 
(4): 165-196. 

-, 1972c. Some subh'ttoral ascidians in More- 



ton Bay, Queensland, Mem. Mus. Qd. 16 (2): 
233-260. 

1972d. Notes on some ascidians from Port 



Jackson, Botany Bay and Port Hacking. Proc. 
Linn. Soc. N.S.W. 97 (4): 241-257. 

1972e. Fauna of the Gulf of 



Carpentaria: 2. Ascidiacea (Chordata: Tunicata). 
Fish. Notes N.S. 3 (3): 39-57. 

1974. The evolution and distribution of 



Australian tropical Ascidiacea in Proc. Second 
International Symposium on Coral Reefs Vol. 1, 
p. 405-432. Great Barrier Reef Committee, Bris- 
bane. 

Linnaeus, C, 1767. Systema naturae. 1: 1087. 

MacDonald, J. D. 1859. On Ihe anatomical characters 
of a remarkable form of compound Tunicata. 
Trans. Linn. Soc. Lond. 22: 373-375. 

Michaelsen, W., 1904. Revision der composition 
Slyeliden oder Polyzoinen. Jh. hamb. wiss. Anst. 
21 (2): 1-124. 

, 1918. Expedition S.M. SchiiT v Pola' in das 

Rote Meer nordliche und siidliche halfte 1895/ 
96-1897/98 zoologische ergebnisse. xxxii. Ascidia 
Pytobranchia und Dictyobranchia des Roten 
Meeres. Denkschr. Akad. Wiss., Wien. 95: 1-120. 
1920. Die krikobranchen ascidien des west- 



lichen Indischen ozeans: Didemniden. Jb. hamb. 
wiss. Anst. 37: 1-74. 

1921, Die Botrylliden und Didemniden der 
Nordsee und der zur Ostsee fiihrended meeres- 
geibiete. Wiss. Meeresunters.^ 97-124. 

1924. Ascidae Krikobranchiae von Neu- 



Spencer Gulf, St. Vincent Gulf and Encounter 
Bay. Trans. R. Soc. S. A. 96 (1): 1-52. 



seeland, den Chatham und den Auckland Inseln. 

(Papers from Dr Th. Mortensen's Pacific Expe- 
dition 1914-16). No. XXII. Vidensk. Meddr 

dansk naturh. Foren. 77: 263-434. 
, 1927. Einige neue westaustralische ptycho- 

branchiate asciden. Zool. Anz. Leipsig. 71: 193- 

203. 
, 1930. Ascidae krikobranchiae. Fauna Sud- 

west-Aust. 5 (7): 463-558. 
Michaelsen, W. and Hartmeyer, R., 1928. Ascidiae 

diktyobranchiae und ptychobranchiae. Fauna 

Siidwest-Aust. 5: 251-460. 
Millar, R. H., 1953. On a collection of ascidians 

from the Gold Coast. Proc. Zool. Soc. Lond. 123 

(2): 277-325. 
, 1955. On a collection of ascidians from 

South Africa. Proc. Zool. Soc. Lond. 125 (1): 

169-221. 

, 1960. Ascidiacea. 'Discovery' Rep. 30: 1-60. 

, 1963. Australian ascidians in the British 

Museum (Natural History). Proc. zool. Soc. 

Lond. 141 (4): 689-746. 
, 1966. Ascidacea. Port Phillip Survey. Mem. 

natn. Mus. Vict. 27: 357-375. 
Monniot, F., 1969. Sur une collection d'ascidies 

composees de Dakar. Mem. Mus. natn. Hist. 

nat., Paris. 41 (2): 426-457. 
Monniot, C. and Monniot, F., 1974. Ascidies des 

lies Kerguelen recoltees par P. M. Arnaud. 

Tethys 5 (4): 715-734. 
Oka, A., 1935. Report of the biological survey of 



ASCIDIAN FAUNA OF WESTERN PORT 



95 



Mutsu Bay. 28. Ascidiae Simplices. Contribution 

from the Marine Biological Station Asamuchi. 
Qtjoy, J. and Gaimard, P., 1834. Voyages de de- 

couvertes de l'Astrolabe 1826-29. 'Mollusques.' 

Zoologie 3: 559-626; 4: 304-306. 
Redikorzev, V., 1927. Zehn neue ascidien aus dem 

Fernen Osten. Zool. Jb. (1) 53: 373-404. 
Ritter, W. E. and Forsyth, R. A., 1917 Ascidians 

of the littoral zone of southern California. Univ. 

Calif. Pubis Zool. 16: 439-512. 
Rowe, F, W. E., 1966. A review of the genus Diplo- 

soma Macdonald, 1859 (Ascidiacea: Didem- 

nidae) with a description of the proposed neotype 

of Diplosoma listerianum (Milne Edwards), 

1841. Ann. Mag. nat. hist. (13) 9: 457-467. 
Savigny, J. C, 1816. Memoires sur les animaux sans 

vertebres. Pt. 2: 1-239. Paris. 
Sluiter, C. P., 1890. Ascidiae simplices. Naturk. 

Tijdschr. Ned. Indie. 50: 329-348. 
, 1895. Tunicaten. In Semon, R. Zoolo- 

gische forschungreisen in Australien und den 

Malagischen Archipel. 163-166 Denkschr. med.- 

naturw. Ges. Jena 8: 163-166. 

1898. Beitrage zur kenntniss dei fauna von 



Sudafrica II Tunicaten. Zool. Jb. (Systematik) 
11: 1-64. 

-, 1900. Tunicaten aus dem Stillen Ozean. 



Zool. Jahrb. Syst. 13: 1-35. 

-, 1904. Die tunicaten der Siboga Expedition. 



Pt. I. Die socialen und holosomen Ascidien. 
Siboga Exped. 56 A: 1-26. 

-, 1909. Die tunicaten der Siboga Expedition. 



Pt. 2. Die merosomen ascidien. Siboga Exped. 
56B: 1-112. 
Sollas, I. B., 1903. On Hypurgon skeati, a new genus 



and species of compound ascidians. Quart. /. 

Micr. Set. (n.s.) 46: 729-735. 
Stimpson, W., 1852. Several new ascidians from the 

coast of the United States. Proc. Boston Soc. 

nat. Hist. 4: 228-232. 
Tokioka, T., 1949a. Contributions to the Japanese 

ascidian fauna 1. Ascidians collected by Prof. 

Mijadi and Mr Masui during the bottom survey 

1939-40. Pubis Seto mar. biol. Lab. 1: 1-18. 
, 1949b. Contribution to Japanese ascidian 

fauna II. Notes on some ascidians collected 

chiefly along the coast of the Kii Peninsula. 

Pubis Seto mar. biol. Lab. 1 (2): 39-65. 

1950. Ascidians from the Palao Islands. 



Pubis Seto mar. biol. Lab. 1 (3): 115-152. 

1952. Ascidians collected bv Messrs Renzi 



Wada and Seizi Wada from the pearl oyster bed 
in the Arafura Sea in 1940. Pubis Seto mar. biol. 
Lab. 2 (2): 91-142. 

1953. Ascidians of Sagami Bay. (Iwanami 



Shoten), Tokyo. 

, 1958. Contributions to Japanese ascidian 

fauna XII Sporadic memoranda. Pubis Seto mar. 
biol. Lab. 6 (3): 313-325. 

1967. Pacific Tunicata of the United States 



National Museum. Smithsonian Institution Bulle- 
tin 251: 1-247. 

Van Name, W. G., 1902. The ascidians of the Ber- 
muda Islands. Trans. Conn. Acad. Arts Sci. 11: 
325-412. 

, 1921. Ascidians of the West Indian region 

and south-eastern United States. Bull. Amer. 
Mus. nat. Hist. 44: 283-494. 

1945. The North and South American 



ascidians. Bull. Am. Mus. nat. Hist. 84: 1-476. 



MORTUARY CUSTOMS OF NORTHEAST ARNHEM LAND: AN 
ACCOUNT COMPILED FROM DONALD THOMSON'S FIELDNOTES 

By Nicolas Peterson 1 

Dept. of Prehistory & Anthropology, Australian National University. 

Tf a man could but follow all that takes place when a yarkomrri [important] man dies he 
would understand almost all of the culture of these people.' — Fieldnotes, July 29, 1937. 



On learning of a man's death close female 
relatives throw themselves on the ground and 
hit their heads with knives, bone points or 
sticks, until blood flows. Some close male rela- 
tives may weep and the son of the dead man is 
likely to become angry and aggressive towards 
his father's enemies. Like the son, the actual 
sister's son may also become angry, recalling 
past quarrels in which his mother's brother had 
figured, regardless of whether he had been in 
the right or wrong. Distant male relatives in the 
camp at the time of death sit quietly with 
bowed heads. 

The wife of a deceased man usually sits 
down beside him, places the head in her lap 
and with her left arm around the body cries 
all day. At night she may lie beside him sur- 
rounded by other camp members who weep and 
sing through the night. The songs indicate the 
path the deceased's spirit, birrimbir, should 
take. 

If people bring news of a death to a camp 
they do not announce the name of the person 
but only mention that somebody has died. A 
senior man in the group then sings a song 
formally announcing the death and at the end 
of the song identifies the person and indicates 
the cause of death without using their name. 
BODY PAINTING 
Most deceased people are painted with a clan 
design, mlntjL The design painted on the body 
should be and usually is that belonging to the 
person's own clan. However, absence of the 
right people may mean that the clan design of 
the actual MM clan is used. Once painted the 
design must not be seen by women or children 
so the painting is often carried out at the edge 
of the camp and when the painting is completed 
the corpse is covered with paperbark, only the 
face being left bare. Young men who see a clan 



design for the first time have underarm sweat, 
bunggan wurdoi, of an older man, rubbed over 
their eyes. 

The body is first rubbed with red ochre and 
then painted by one or two men, preferably 
of the opposite moiety to the deceased, who 
are good hands at painting, kong mintjimirri 
The most frequently chosen relatives are from 
the categories FZS, MBS, ZS, MB and MF/ 
FMB. If the painters are of the same moiety 
they are likely to be close WMB, ZDS, or FZDS 
but never actual F, B or S. Whoever they are, 
they are referred to as the kong wukundi, 2 
hands tabu from death, and after they have 
completed the painting coat their arms and 
hands with red ochre, eat apart for about a 
week, refrain from sexual intercourse and do 
not go near water. They lived with their wives 
some distance from others in the camp. The 
kong wukundi and their wives cook all their 
food in a sand sculpture (wandjur — discussed 
below) and put all their food scraps into 
another. 

Case 1. Bodv painting (see Plate 4). An old Obul- 
karra [Wulkara of Warner, see 1958: 46] woman 
died at Milingimbi on Sept. 19, 1935. She had 
been declining for some months and was very thin 
from the effects of leprosy. As she was old there 
was very little crying, but Thomson was suprised 
at the apparent indifference of all the imme- 
diate relatives. The day following the death a 
Wunguri clansman who called the deceased muk- 
kulmal (FZ) and a Tjambarapoingo man who 
called her momalkor (WMM) painted the body. 
The painting was carried out under the shade 
of a big tree about fifty yards from the camp. The 
husband and a few other men came to and from 
the place at which the body was painted from 
time to time. Ordinarily her husband would have 
assisted in the painting but he was too old and 
could not see properly. The brothers of the de- 
ceased, as is always the case, could not touch the 
body. At the conclusion of the painting the kong 
wukundi painted their hands and arms below the 
elbow with red ochre and that night held a mant- 
jarr ceremony. This took place on the fringe of 



97 



98 



NICOLAS PETERSON 



the camp. A sand sculpture, wandjur (see below) 
representing the clan well of the deceased was 
moulded on the ground. The sculpture was associa- 
ted with the edible corm of Eleocharis dulcis, called 
rakai, but the significance of this was hard for 
Thomson to follow at the time [as he had only 
just arrived in Arnhem Land]. The men sat around 
the sculpture singing for an hour or two, and then 
the women danced behind them. A fire was lit 
in the centre (manotji [literally eye but also used 
of a certain kind of well]) of the sand sculpture. 
Leaves were then heated in the fire by a man 
whose mother came from the 'Wulkarra [that is 
Obulkarra] side'. All the immediate relatives gath- 
ered inside the sculpture about the manotji, and 
a 'big' man dalkarramirri, called out in loud voice: 
'KuritaF (fire). 

'Ye'h- replied the participants, with a long his- 
sing shout. 
Fire was called because the ancestral woman, 
mialk kortjurino, burned the grass to clear the 
ground while looking for rakai. 

The participants then shouted 'Wap wap! wap!', 
the sound of the fire burning. 

The dalkarramirri then called out a series of 
big names with the participants answering 'Ye'h!' 
to each. 

'NunimarraV (big name [of fire?]). 
'Baltjau'wuma' (a big light made by fire flaring 

up in thick grass). 
'Matauwupuptum' (leaping tongues of fire). 
'Birraudun' (cleansed by burning). 
Throughout the calling leaves were being heated 
in the fire and used to strike the bodies of the 
people standing in the sculpture. After use they 
were burnt in the fire and to the accompaniment 
of singing, smothered with earth, 'that mialk 
[woman] walk about now — come along clean place'. 
Then the men sang of marratnatta, a rodent 
that followed the fire and established itself in the 
plains after the burning; and then of dog (workan) 
who smelt the rats; and after that of plain cock- 
atoo (Corella sp.) kai karra. 

The ancestral woman now looked about for 
the rakai. Next the men sang of mist (kardany) 
which is like fine smoke; then of a spider making 
its [nest/webb?] in the damp grass. The last song 
was about the wind that follows the time of burn- 
ing the countryside. 

Close relatives are not free from all tabu until 
after a second cleansing ceremony. 

HAIR 

Before burial of the body all the head hair 
is pulled out by the kong wukundi. The beard 
is also pulled out with the aid of hot bees wax. 
The hair is kept in a basket and sometime 
during the following weeks is sent by the kong 
wukundi to a fairly distant relative of either 
moiety to turn into a string decorated with 
feathers, marngarai (Kopapoingo, Tjambara- 
poingo and Koiyamillilli) or yiritpal (Wunguri). 
The maker of the yiritpal or marngarai rubs 
red ochre on the hair as soon as he receives it 




MEM. NAT. MUS. VICT. 37 PLATE 4 




An Obulkarra woman being painted with a clan 
design after death (Case 1). 



MORTUARY CUSTOMS 



99 



and is given a present of vegetable food from 
the marramorkoimirri (the deceased's patri- 
clansmen). Usually the maker is of the opposite 
moiety to the deceased but he does not have to 
be; common choices are people in the category 
of MF/FMB, MB, MBS, WMMB. One to two 
years later it will be completed and returned 
to the close kinsmen of the deceased eventually 
being given to his son, if adult. A large presen- 
tation is made to the maker; this formerly 
included vegetable food, cycad bread and 
hooked and short spears. The string is then 
used as a belt to be worn in ngarra ceremonies 
and during fights and makaratta peace-making 
settlements. Eventually the string is cut up to 
form the 'arms' of men's sacred baskets. 

BURIAL 

Either the same day as the painting or the 
day afterwards the body is buried. There are 
two types of burials: either in the ground or 
in a tree. 

A grave, molo (referring specifically to the 
heap of earth covering the grave) is usually 
1 m deep and long enough for the corpse to 
lie extended, on its front. There seems to be 
some variation of opinion as to which way a 
deceased man's head should point. In a dis- 
cussion on the matter a Djinang and a Lia- 
gallauwumirri man maintained it should be 
towards the clan well while a Kopapoingo and 
Tjambarapoingo man maintained it should be 
eastwards. 

If a body in a grave is covered with heavy 
logs and stones it usually signifies that it is 
to be left for good and there is no one to carry 
the bones about so that the bones will not be 
removed. There is no belief as to any ill result 
to the spirit from this practice. Usually, how- 
ever, the bones are expected to be dug up. 
In this case the body is covered by a sheet of 
paperbark, the earth replaced and poles and 
stones placed on top to stop dingoes, lizards 
or dogs eating the flesh. If a camp dog does eat 
the flesh it becomes wukundi, and may be 
killed. If it is not killed it will be put through 
a cleansing ceremony and any food that the 
dog catches before the cleansing can only be 
eaten by a male owner of the dog. Even after 



the restriction has been removed the wife of 
the owner or other women will have to make 
a ceremonial presentation of food to the marra- 
morkoimirri as soon as she eats food the dog 
has killed. 

A grave may be located in one of three 
places. Where the deceased is a child, it is 
often buried in the camp of the parents who 
sleep beside the grave until exhumation. If it 
is an adult the body is usually buried outside 
the camp, but it may may be in it if the people 
plan to abandon camp immediately. 

In the southeastern part of the Murngin area 
around Blue Mud Bay there is the third kind 
of location: the collective burial ground. 

Case 2. Visit to communal burial ground. On 
October 18, 1935, Thomson visited a communal 
burial ground or wukundi place at Blue Mud Bay. 
It was situated on the edge of Marrakulo territory 
at a place called Mange'yall, 5 km from the 
Aborigines' camp. [Warner (1958: 49) refers to 
the people of this area as Marungun and notes 
that their Vaterhole' and 'country' are called 
Mangaia which he glosses as 'stench of a dead 
man' .J 

A Marmariny man had died some 2-3 weeks 
earlier. His body had been taken to this place 
and put on a platform. The platform stood about 
1-50 m high with the body upwards and roughly 
covered in paperbark. The ground around the 
platform was sculpted into a wandjur pattern. The 
body was sharing the platform with a number of 
other bones from skeletons, most of which were 
covered with red ochre. In this case the head of the 
body was placed towards the east so that the malli 
[shade or shadow] could go to Buralko [the land 
of the dead]. 

On this day Thomson visited the burial ground 
in the company of five men: Raiwalla, his Mild- 
jingi companion; Taudauongo, the actual elder 
brother of the deceased; an old Ritarango man; 
Djimbaron, a Dai'i man; and Marrilyanwi, a 
Marmariny clansman who called the deceased son 
and had been one of the kong wukundi. The 
other kong wukundi were Liawulpul a Bidingal 
man who called the deceased du'wai (FZS) and 
Marakuri a korrong of the deceased (FZDS). 

The party set out from the camp travelling 
across salt pans. A line of fires was burning in 
the short grass between the camp and the burial 
ground. As the grass was sparse and short and 
provided neither food nor cover for the animals 
this was surprising. On enquiry it was found that 
the fires had been lit to 'block in wukundi — 
might be smell go all round', i.e. to cut it off from 
the camp to which it is believed to be more or 
less connected by smell. When the party was 
within 0*50 km of the place the Aborigines re- 
quested Thomson to leave the water he was 
carrying lest the malli [shade] of the wurkaidi 
(larvae ancestor) might 'go into it and make 



100 



NICOLAS PETERSON 



(him) sick (rerri) . . .'. They translated the term 
rerri, generally used for sickness of any kind, as 
leprosy. 

None of the Aborigines carried spears or spear- 
throwers which was most unusual; this was said 
to be one way of helping to avoid sickness. As 
they neared the spot, Marrilyanwi took charge. 
Thomson was told not to stay too long or go 
too close lest he should fall sick. They all ap- 
proached the platform by a roundabout path to 
take them upwind. They conversed in whispers 
and walked slowly with the arms folded to avoid 
being 'flash'. The little outcrops of stone that 
appeared on the flat salt pans were carefully 
avoided. A couple of hundred yards off they halted 
and Marrilyanwi rubbed his hands under his 
armpits and then over Raiwalla's arms and legs 
before kneeling down and biting his knees and 
shins all the way down. Marrilyanwi then spat or 
hissed in the direction of the wukundi place. He 
did the same to Thomson. When they moved off 
Raiwalla was told to walk behind Marrilyanwi in 
his footsteps. When they reached the platform they 
looked briefly and then moved away and turned 
their backs while Marrilyanwi approached the plat- 
form and smashed a pipe close to it in order to 
pacify the malli so that it would not follow them. 
'Chuckit smoke along him, that's all', explained 
Marrilyanwi. They left and the Aborigines washed 
in a salt pan a few hundred yards off. 'Wash'em 
sweat, ground, that maggot him bite you and me— 
malli — you and me no been see that malli\ one of 
them commented. 

On October 24, Thomson visited the burial 
ground again with a Ritarango man, Wuruwul, 
who had not been there before. Raiwalla and 
Thomson did not have to go through any of the 
procedures of the previous visit, although they did 
approach upwind again. Wuruwul, a stranger to 
the place, was very fearful of sickness, particu- 
larly because he was somewhat fat. As a precaution 
against sickness he had his knees and elbows bitten 
(see Plate 5) and left the area well before the 
rest of the party. 

The alternative to ground burial is exposure 
on a platform, either built in a tree and called 
djamba or free standing like the kind used in 
house building and called katauwurro. [Warner 
states (1958: 433) that the body on a platform 
is placed face up so that when the adominal 
wall breaks the intestines will not fall.] Thomson 
found that although this was true for the eastern 
half of the area the people to the west of the 
Ritarango place the body on its front. 

The choice of burial mode depended on 
several factors. Small children and old people 
were usually put in the ground and active people 
in their prime, male or female, were placed 
on platforms where the flesh dries more quickly 
and the bones become cleaner. If a person were 
killed in a miringo raid they were usually put 



in a tree by their relatives so that the people 
could leave the area immediately. 

There is a third mode of disposal found 
among the Burara who Thomson reports as 
eating young men, women and children, after 
roasting in ashes [although Thomson did visit 
the Burara it is not clear how much of the 
following information was obtained by talking 
with and observing Burara people and how 
much was supplied by their Glyde River neigh- 
bours who hold the Burara in low esteem]. 

The dead are eaten by all relatives with the 
exception of M and MB because the 'two fella 
been carrim along bindji [belly]'. Bodies to be 
eaten have an incision made in their left side 
through which the viscera are removed. The 
liver is eaten but the heart, penis and vulva 
are dried and carried in a special small basket 
(pulupur in Burara) or in a matjitji to increase 
hunting effectiveness. The lungs and stomach 
are buried. 

Because the body is eaten the western Burara 
do not paint it, although the eastern Burara, 
under the influence of their western neighbours, 
the Wallamango and Yarnango, do. These latter 
groups rarely eat their dead but do inspect the 
internal organs, for signs of sorcery such as 
sores (tjitjt) on the kidney, heart or liver, by 
making an incision between the crest of the 
ilium and the last rib on the left side. Now that 
there are marngit medicine men in the area — 
a new tradition from the south [see Thomson 
1961] — the people no longer perform this kind 
of investigation. 

POSSESSIONS 

The Djinang speaking peoples pull down the 
deceased's hut and eventually set fire to it when 
the bones have been exhumed. The main pos- 
sessions of a person, such as his spears or a 
canoe he has made, are treated throughout the 
area in the same way as people. They are sym- 
bolically cleansed at a mantjarr ceremony and 
in the case of canoes rubbed with red ochre 
often on two separate occasions. If a man has 
been speared, his possessions are broken and 
pieces given to his relatives in camp which an 
informant interpreted as 'that mean I push all 
these people go for fight'. Each piece of broken 
possession used in this way is called maidjaballa. 



MEM. NAT. MUS. VICT. 37 PLATE 5 




m 




Wuruwul having his elbow bitten as a precaution 
against sickness prior to visiting the communal 
burial ground in Blue Mud Bay (Case 2). 



MORTUARY CUSTOMS 



101 



If a person does not want to fight he makes 
a new dilly bag and gives it back to the man 
(usually the elder brother of the deceased), 
who distributed the maidjaballa. The broken 
pieces are given most frequently to relations 
in the categories MF/FMB, MMB, FZS, 
MBS, ZS. 

If a person dies from a cause other than 
spearing his possessions are also broken up and 
distributed to various other residential groups 
where there are close relations of the deceased. 
These possessions are then used in the clean- 
sing ceremonies. 

Thomson reports a case where following the 
death of an important man a restriction was 
placed over a large area of land. 

Case 3. An area of land placed under restriction. 
While at Katji [on the mainland south of Milin- 
gimbi] camp in January 1937 Thomson noticed 
red blazes on the trees along the path to Derby 
Creek. These were to free the area about the Katji 
River from a restriction that had been imposed 
at the death of an important man. 

The restriction had been imposed at the death 
of Raiwalla's father-in-law [Raiwalla, a Mildjingi 
clansman, was Thomson's guide and friend] be- 
cause of his influence when alive and because 
he had spent much of his life in that area. The 
restriction was not removed until after the bukubut 
[exhumation ceremony], when the trees were 
painted and a lire lit to burn off the grass and 
cleanse the area- — both literally and figuratively. 
The fire was started by burning his old camp 
with a fire of ironwood. After the burning off the 
women went out and collected root foods in the 
area and brought them back to the son and bro- 
thers of the deceased. If any outsider eats food 
from the area while the country is under the 
restriction the close relatives resent this and try 
to kill the offender by sorcery or with an actual 
war party (miringo). Such restrictions do not apply 
in remote areas but only when important people 
{yarkomirri) die in the vicinity of an important 
camping place. The death of the wife or daughter 
of an influential man can also lead to the same 
restriction Tjambarapoingo, Kopapoingo and 
Ritarango peoples have the same custom. 
CLEANSING CEREMONIES AND GROUNDS 
Cleansing ceremonies of the kind described 
in Case 1 are held at several stages following 
death. After burial the clan song cycle of the 
deceased is sung about a circular sand sculpture 
and all men, women and children present, 
together with the larger possessions are dusted 
with heated leaves (mantjarr) to drive the nialli 
(shade) of the deceased away, to render the 
hunting weapons effective and the other pos- 



sessions safe to handle. 

At a second ceremony of similar form the 
participants throw pieces of the deceased's 
possessions into the fire burning in the small 
depression which forms the focus of all the 
cleansing ceremonies at this stage. At this 
second stage the women usually dance while 
the men are singing the clan cycle. 

A week or two later the third cleansing 
ceremony, called bukulup (forehead-washing), 
is held. The small circular sand sculpture used 
in the previous ceremonies now gives way to 
a much larger and more elaborate representa- 
tion of the clan well. Starting in the early hours 
of the morning the men sing the clan songs 
and then once the sun is up the close relatives, 
male and female and the kong wukundi, wash 
standing in the well and rub red ochre over 
themselves. This ritual frees the kong wukundi 
from all tabus. The patriclansmen make a pre- 
sentation of food to members of the opposite 
moiety. 

The sand sculptures are also used in two 
other contexts. Most frequently in the curing 
of sores or wounds but also around graves and 
burial platforms (see Case 2). Several clans 
may share the use of a particular design and 
a single clan may have several designs relating 
to different places with differing degrees of 
importance. Figure 1 shows sketches of six 
grounds seen in use by Thomson. 

EXHUMATION AND FLESH DISPOSAL 
After a month or two the bones are exhumed. 
The men who perform this task are also known 
as kong wukundi [possibly also as kong djok] 
but are not necessarily the same people who 
buried the body. They may be of either moiety, 
with the reservation that a man may not assist 
in the exhumation of his siblings. 

The grave is usually dug out either with bare 
hands or with a sharpened stick. Among the 
Djinang speaking groups only males are present 
at the grave. One will sing and another play 
the digeridoo. Among the people to the east, 
women are present at the exhumation and dance 
while the men sing. 

The treatment of the flesh varies with the 
kind of burial and the area. Among the peoples 



102 



NICOLAS PETERSON 




SOME WANDJUR SAND SCULPTURE DESIGNS 

(Running from left to right and top to bottom) 

1. Marango clan (dua moiety) wandjur representing 
the bee hive yarrpain [referred to as 'long-nose 
sugar bag' in Aboriginal English on account of the 
relatively long entrance tunnel]. The central 
circle represents the eye of the clan well; the 
rectangle surrounding the well and the area im- 
mediately above it is referred to by the sacred 
name bambula. The small rectangle had a pole 
483 mm tall erected in it, called warrinman, repre- 
senting an ancestral hero. The ground was used 
on July 24, 1937, at Milingimbi for the cleansing 
of two small girls. The same design may also 
be used by Tjambarapoingo speaking clans. 

2. Birkilla clan (yiritja moiety) wandjur representing 
the bee hive birkurda at a place called Yarrakka 
in Arnhem Bay. The small circle at the end rep- 
resents the entrance (ngorro — nose) of the hive. 
[This wandjur should be compared with the illustra- 
tion published by Thomson in the 'Illustrated 
London News' for February 25, 1939, page 294,] 
This ground is the most elaborate form of the 
wandjur and only used for important men; others 
have a simplified, but recognizably similar, ver- 
sion. The ground was used on August 7, 1937, 
to cure a Birkilla/Kopapoingo man with a sore. 

3. Birkilli clan {yiritja moiety) wandjur representing 
waitjura [? a fish]. The piles of white sand are 
sores (tjitji or mapan) made by a crab (mirriya 
or katjirri) . The location referred to is Karraparra 
in Blue Mud Bay and the design is also used 



by Yituwa clansmen of that area. The ground was 
used on August 7, 1937. 

4. A dua moiety wandjur used by Liagauwumirri. 
Maiyarrmaiyarr and other Tjambarapoingo speak- 
ing groups united by the track of the Djanggauwo 
sisters. The ground represents springs left by the 
sisters whether they thrust the 'yam' sticks into the 
ground. The springs are called milmindjarrk [and 
are marked by being freshwater sources in areas 
of salt surface water. The arrows appear to indi- 
cate the direction of flow of the waters beneath 
the wells]. The ground was used on July 30-31, 
1937, at Milingimbi for a Maiyarrmaiyarr man 
who was drowned when a canoe turned over 
during a storm in the Cadell Straits. There were 
six people in the canoe: two men escaped but 
a blind man and a second man [it is not clear 
which was the Maiyarrmaiyarr man] with his son 
and daughter were drowned because they were 
encumbered with turtle hunting gear. The ground 
measured 15 m overall. 

5. Kolumalla clan (dua moiety) wandjur represent- 
ing mar'ndi [?] The ground was outlined in white 
sand and used on August 14th, 1937. 

6. Warramirri clan (yiritja moiety) wandjur repre- 
senting a whale, woimirri. The small circle is the 
rectum above the tail. The central rectangle is both 
the whale's stomach and the manotji or eye of 
of the clan well. The soil forming the outline was 
raised up 100-125 mm and whitened with sand. 
This ground was used on August 14 and 15, 
1937, to cure a child of the Wunguri clan who 
had sores. The child's full MMB came from the 
Warramirri clan. 



MORTUARY CUSTOMS 



103 



east of the Glyde River, including the Kanal- 
pingo and Djinba, flesh from a grave burial 
is put back into the ground. Flesh from a plat- 
form burial is placed in paperbark and left in 
a forked tree nearby to be destroyed naturally. 
The platform itself is pulled down and buried 
in the sand sculpture in which it was standing. 
Among the Djinang, Mildjingi, Balambi, 
Wullaki, Burara and all groups westward of 
the Glyde, the flesh is kept and at the bukubut 
ceremony placed in a special hollow log called 
larkan djammurmur. 

Case 4. Larkan djammurmur form of flesh dis- 
posal. Early in the morning of November 11, 1936, 
the Wullaki group at Katji started to sing in 
preparation for a bukubut. The Wullaki people 
were joined by some Milli'ereng clansmen because 
the dead woman's mother was of this clan. 

The body had been buried in the ground. A few 
days earlier it had been exhumed and the flesh 
roughly stripped from the body and wrapped in a 
paperbark bundle. The bones were in a second 
bundle. The men had then cut a tree for the 
larkan [coffin] and the women collected cycad 
nuts for a food presentation. While the cycad 
nuts were being leached the larkan was fashioned 
and a marraidjirri meri (an effigy of an ancestral 
spirit) made. The preparation of the larkan in- 
cluded singeing it, cutting the spikes on one end, 
and painting it. On the morning of November 11, 
1936, the men were seated in the shade of a 
clump of trees some distance from the camp with 
the larkan. The bundle of flesh was apparently 
[this not unequivocally clear in the notes] already 
inside the coffin. The wrapped bones and a mar- 
raidjirri meri called Kanangalkngalk were also 
nearby. This ancestral spirit was responsible for 
the people using a log coffin. 

An informant explained that in the distant past 
(millegidji) there were spirits (meri or morkoi) 
that were neither animals nor men and who still 
live in the bush today. These spirits were never 
men but a race of their own. One of these spirits, 
Kanangalkngalk, is still alive today and some Wul- 
laki people even claim to have seen him in the 
monsoon forest. Kanangalkngalk has two wives 
and some children, none of whom is a threat to 
the people like the spirits of deceased human 
beings. The marraidjirri meri represent Kanan- 
galkngalk. 

They say that in the distant past Kanangalkngalk 
cut down a hollow tree. The tree fell and as it 
fell water started to pour out of it. He tried to 
hold onto the log as the water flowed out but his 
fingers slipped and the log moved off like a fish. 
The log cut the ground as it went allowing the 
water to flow. Along the way the log heard a 
buralla [publicly used bull-roarer) sing out. Then, 
perhaps because the water told it to, the log 
went underground at Katji carrying earth and 
water with it as it went. The log wanted to go 
down towards the sea but found the ground too 
hard so it came back and let the water go. From 
the end of the Katji lagoons, where 



he turned back, there is no deep water but only 
transient flood waters at the end of the wet season. 
The log came back to Katji and decided to stay 
where the deep pool is beside the camp. At that 
place he gave himself a name: the log said, 1 am 
djammurmur larkan." The people today reflect on 
the fact that they take fish, water snakes {Hypsir- 
hina) and wild taro from Katji where the larkan 
walked around. That is why they paint them on 
the log coffin and cut the long 'fingers' into the 
mouth of the log, representing the jagged end 
which resulted from it breaking the ground and 
making Katji River. 

The men around the larkan began to sing a 
song about the wullawarri fish drawn on the log. 
Then they picked up the log and danced with it, 
making short lunges and rushes, replacing it on 
the ground at the end of each movement. (See 
Plate 6.) The log was then erected on the open 
ground near the camp and the marraidjirri and 
bones were carried toward the camp by two old 
men. The dance that followed was called after the 
spirit, Kanangalkngalk meri. The main body of 
men danced forwards looking for the meri carried 
by an old man, who represented the male spirit. 
The one who carried the bones represented a 
female morkoi [spirit] [Kanagalkngalk's wife?]. 
The first old man kept dancing forward with the 
meri to reassure himself that the other man had 
the bones. 

The women danced their slight shuffling dance 
from one foot to another on the fringe of the 
dance area. The dance concluded with an old man, 
dalkargrining [dalkarramirri] calling the big names 
of the maraiin of the deceased and of her country. 
Then the bones were handed over to the actual 
younger sister of the woman's mother, the real 
mother having died. As she received the bones in 
their bundle the marraidjirri meri was placed on 
top and she walked off with the whole lot. 

At a later date the string on the marraidjirri 
meri is removed and made into ngaimbak [arm 
bands] and used in decorating a bati giwillir [a 
kind of men's basket]. This basket is then pre- 
sented to the man who made the string. The con- 
clusion to the bukubut comes sometime later. A 
presentation of food is made to the kong wukundi 
by the close relatives of the deceased, usually the 
F. duwe (either FZS or D), MB and EB if it is 
a man that has died but not in the case of a 
woman. 

This food is not eaten bv the full F or MB 
but FEB and FYB, duwe and kalli (MBS/D) do 
share in it. 

In the past the Djinang people removed the 
flesh from the buttocks, washing it free of the 
soft and more putrid surrounding flesh and tied 
it up in paperbark. Later these parcels of flesh 
were cooked outside the camp, wrapped in 
grass and paperbark and hung around the neck 
to increase hunting effectiveness. Some people 
would go a step further and soak the flesh in 
a mixture of honey and water and then nibble 
a fraction with their eyes closed. 



104 



NICOLAS PETERSON 



The bones are washed and wrapped in paper- 
bark, of if the flesh is not entirely removed, 
left exposed in a forked tree. During the 1-2 
weeks before the bukabut ceremony the bones 
remain outside the camp and may not be seen 
by the women. 

The kong wukundi camp apart for several 
days. The wetter the body the longer the period 
of restriction. If it is particularly sloppy the 
men eat with a bone point (pringal) or any 
sharpened stick because the fluids will have 
penetrated their finger nails making them smell 
for some time. 

A day or two after the exhumation the kong 
wukundi participate in a cleansing ceremony 
singing all night and washing in the morning in 
a sand sculpture. After washing they cover 
themselves in red ochre. Several days later they 
have a clan design painted on them which re- 
leases them from all restrictions. 

Case 5. Exhumation. On January 13, 1937, the 
bones of a Djinang man named Lamieri, dua 
moiety, buried at Gillere in Millierieng territory 
were exhumed fsee Plate 7). 

Only a few people went to the area of the grave 
where the man had been buried 6-7 weeks before. 
Those not directly involved in exhumation stood 
upwind of the grave. 

Two men were involved in handling the bones. 
The man who removed the bones was the adopted 
father of the deceased from the same country 
[i.e. clan] named Balambarri. He was assisted bv 
Makani a Mildjingi man a 'ZS 1 of the deceased, 
who was married to two of his daughters. 

On the way to the grave there was some discus- 
sion as to whether the body was soft enough 
yet to make the removal of the bones easy. The 
grave itself was unmarked except for a plain cir- 
cular sand sculpture near the head of the grave 
and a heap of wood lying on top to keep the dogs 
off. The body was about 1 m below the surface, 
lying face down on a layer of grass and completely 
extended. The grave itself was in loose, well- 
drained sandy soil about 90 m from a creek. The 
soil was removed largely with bare hands but use 
was made of a canoe paddle that happened to have 
been in the camp and brought along. 

An old clansman of the deceased, a classi'ficatory 
F, sang_ to the accompaniment of clapsticks. In 
Kopapoingo and Tjambarapoingo ceremonies 
women are present and dance during the exhuma- 
tion but not among the Djinang. The kong wu- 
kundi examined the body to see that the flesh 
was sufficiently decomposed and finding it was, the 
assistant, Makani, went off to get some water in 
a paperbark trough. 

The deceased had been an old man of not much 
standing so he had not had a clan design painted 
on his chest. 

The adopted father removed the bones, starting 



from the feet and working upwards. Makani poured 
water over the bones as the first man washed them 
thus reducing the period he would be wukundi 
[tabu]. The head was picked up last and washed 
by pouring water in through the foramen magnum. 
Each bone as it was picked up was placed on a 
sheet of bark beside the grave. When they had all 
been removed the grave was filled in again. The 
two men washed and smeared themselves with 
white paint from head to foot: 'everybody no 
more want to smell*. Red ochre is only used ixi the 
final cleansing. During the night a sand sculpture 
1-60 m in diameter was made at Makanrs camp 
and a ceremonial washing called bukulup carried 
out on the following morning (see Plate 8). About 
sunrise the two kong wukundi washed by pouring 
water over one another's bodies and then smearing 
red ochre all over themselves, their spears and 
spear-throwers and immediate possessions. This 
freed the men and their weapons from tabu. 

It is usual to wait two or three days before 
holding this ceremony, but as the camp was break- 
ing up on the following day it was completed 
straight away. 

MARRAIDJIRRI MESSAGE STRINGS 
Marraidjirri is the general term for a class 
of decorated strings whose most common use 
is in the mustering of people for exhumation 
and final disposal ceremonies (see Plate 9). 
Marraidjirri strings differ from mamgarai 
strings in that although many of them incor- 
porate hair of a dead person they are largely 
made of fibre string and are used in a different 
way, (for marngarai strings see under 'Hair' 
above). 

Each clan has its own marraidjirri forms 
representing totems associated with the clan 
(see Table 1). Generally there are several 
forms of the string-like marraidjirri which are 
classed together as bogongo and spoken of as 
'small' in contrast to the elaborate figure em- 
blems such as that mentioned in Case 4 which 
are referred to as big (yindi). 

Besides being used to gather people for 
mortuary ceremonies they are also associa- 
ted with circumcision ceremonies, the social 
development of children and love magic. 
In circumcision ceremonies they are used to 
gather people. The second usage result from 
the first occasion on which a small child 
picks up any natural object such as grass, 
a shell, fruit or small lizard and gives it to 
to its parents. This object is then tied into a 
small bundle and sent off to an acquaintance 
both geographically and socially distant. This 



MEM. NAT. MUS. VICT. 37 PLATE 6 




«-* * 4f"*- ■*** 






Wullaki men taking the larkan coffin to the camp 
for erection (Case 4). 



MEM. NAT. MUS. VICT, 37 PLATE 7 




The exhumation of a Djinang man (Case 5). 



MEM. NAT. MUS. VICT. 37 PLATE 8 







Ritual washing in a wandjur sand sculpture of 
two of the men who participated in the exhuma- 
tion shown in plate 4 (Case 5). 



MEM. NAT. MUS. VICT. 37 PLATE 9 




On August 20, 1935, this messenger, a man of 
the yiritja moiety arrived at Milingimbi wearing 
a dua moiety marraidjirri message string. He had 



come to call the people to Elcho Island for the 
final disposal of the bones of a Tjamborapoingo 
man. 



MORTUARY CUSTOMS 



105 



person fashions the object into his own clan's 
large marraidjirri and presents it to the child's 
parents in a large public ceremony. The parents 
then make a payment of traditional wealth of 
considerable proportions to the maker of the 
marraidjirri. On completion of the ceremony 
the string is removed from the core about 
which it is bound and made into arm bands or 
used for adorning certain kinds of men's baskets 
( bati mindjalpoi ) . Some marraidjirri strings 
may be used in love magic after they have been 
used in one of the foregoing ceremonies A 
sweetheart or errant wife is believed to be 
impelled to follow the man involved when the 
marraidjirri string is looped over her hands. 

The strings sent out to muster people for 
ceremonies are really representations of big (or 
proper) marraidjirri constructed around wooden 
or sometimes paperbark centres and used at 
the bukubut (see Case 4). These solid emblems 
look like rangga (the secret totemic emblems) 
as an informant observed to Thomson, but may 
be seen by women and children and are said 
to represent the clan's ancestral morkoi or 
spirit being. This morkoi is different from the 
clan's ancestral hero, wangar. However, as with 
the clan totem, the string covering is called 
buy it and is equated with flesh; in its broadest 
sense it just means covering. The core is re- 
garded as maraiin (sacred) and identified with 
the bones of the skeleton. 

TABLE 1 

Some of the small marraidjirri strings 
(bogongo) used by various groups 

CLAN/GROUP 

Wanguri 

Birkilli 



Tjambarapoingo 

Kanalpingo 

Liagallauwumirri 

Mandalpoi 
Ritarango 
Durrilli 
Obulkarra 

Mildjingi 



NAME and DESCRIPTION 

Yorko — a round root called Kalun 
in Kopapoingo (Cissus carnosa) 

Komulo — the great billed heron 

Ku'ak — a small bird 

Yukuwa — the root food Vigna 
vexillata 

Tjarrak — a tern {sterna sp.) 

Kalliwur — a large white lily 

Wititj— Snake 

Ku'ak — a small bird 
Malka — -bee (Trigona sp.) 
Yorko — round root (Cissus 

carnosa) 
Kurungur — a small cloud and also 

the wild bean Ipomea pes-caprae 



BUKUBUT FOLLOWING EXHUMATION OF 
BONES ONLY 

When the food for the bukubut ceremony is 
ready, a week or two after exhumatiom a classi- 
factory ZS goes into the bush and picks up the 
bones. Meanwhile the marramorkoimirri pre- 
pare the ground, knoww by different names to 
different clans: 

Liagallauwumirri call it birlimbil 

Djeranggoikoi „ „ djirrkurul (of 

bulmantji or 
shark) 

Birkilli „ „ yallandu (from 

Bukunda, a 
place) 

Mildjingi „ ,, manitji 

When the 'ZS' carrying the bones, ap- 
proaches the ground just outside the camp 
where the ceremony is to be held, the men 
begin to sing. The marramorkoimirri and a 
great crowd of more distant kinsmen dance 
around with spears poised and jab these at 
the bones in their paperbark wrappings as 
they lie in the sand sculpture. 'Him want spear 
that one, open him — wangar (totemic ances- 
tor) been do.' Not all clans carry out the 
ceremony in this way. The western ones, Lia- 
gallauwumirr, Mildjingi and Djinang only sing. 

The Birkilli, Daigurgur, Ritarango and al- 
lied groups customarily spear the parcel, and 
do the same again when they are holding the 
final disposal ceremonies. They open the paper- 
bark with the spears and then sit down and 
wash the bones before placing them in a new 
wrapping, and hanging them from a forked 
stick standing in the middle of the sand sculp- 
ture. At this stage the women and children 
may not see the bones although they can later 
on when they have been red ochred. 

From the late afternoon onwards through 
the whole night the men sing and complete the 
bukubut ground where the bones will be pre- 
sented to the woman who is to carry them. 
This is usually the actual FZ, EZ, adult D, or 
mother if the deceased is a young child. If 
the bones of an older person are given to a 
mother it is always to a classificatory mother; 
they are never in the custody of a sister 



106 



NICOLAS PETERSON 



although she may handle and carry them. In 
the morning these relatives and the MM will 
dance near the song group which is usually 
composed of EB, YB, F. The bones are then 
handed to the MF/FMB, ZH or MMB. Most 
commonly, it is to the ZH who, then hands 
the bones to the deceased's FZ who will carry 
them during the following weeks. 

Previously the F, S, ZDS and ZS of the 
deceased will have made a bark container, 
tarra, decorated with the deceased's clan design 
for the bones to be carried in. The Mildjingi 
and Liagallauwumirri do [may?] not have this 
custom. There is then a ceremonial present- 
ation of food by all the helpers in the various 
stages of the ceremony to the marramorkoi- 
mirri. 

Often the skull is not placed in with the 
other bones but carried separately by WB or 
another ZH. The bones are carried for 1-5 
weeks. At the end of this period they are again 
hung from a forked stick in camp and only 
moved on shifting camp, until the final disposal 
of the bones in a hollow log coffin ceremony. 

Case 6. A Wullaki hukuhut ceremony. A bnkuhui 
ceremony at which the bones of a dead Wullaki 
speaking man of yiritja moiety were handed over 
to the deceased's sisters was held at Katji on 
October 3 and 4, 1936. 

When Thomson arrived in camp late in the 
afternoon the ceremony was about to begin. The 
bones were hanging in a small shade (kurngan) 
along with the marraidjirri meri. The marraidjirri 
represented a wasp's nest called banal and was 
decorated with a picture of the little green pigeon, 
work'miringo (Chaleophaps chrysochlora). This 
bird is associated with the paper wasp in areas 
of monsoon forest. 

Although the deceased man was Wullaki the 
bukubut was carried out with a Mildjingi song 
sequence because the Wullaki relatives had handed 
the bones to the Mildjingi [Thomson has Raranggal 
malla at this point in his notes, but on the first 
page he equates Mildjingi malla with Raranggal 
malla] clansmen as a friendly compliment. 

The ceremony began with singing to the accom- 
paniment of clapsticks and didgeridoo and con- 
tinued for some hours into the night until every- 
body was supposed to be asleep. Each man then 
seized a torch of lighted paperbark and started to 
dance, first encircling the shade with the bones 
and marraidjirri in it, and then moving into the 
camp crying berk! berk! berkberk ko ye'h ko yeh. 

They encircled the whole camp where everyone 
pretended to remain sleeping and then trotted back 
in single file to the shade. This was the dance of 
the flying fox. The singing then continued. When 
they began the song about the jungle fowl (gulla- 



uwurr) the men started to dig a long serpentine 
path, which eventually measured 42 m, from the 
shade to the point where the men constructed a 
representation of the bird's nest. At intervals the 
men working on the road and the nest cried out 
kurrkun kurrkiin djue wurak — grr'rr in imitation 
of the jungle fowl; these cries were heard inter- 
mittently through the night. Jn making the nest 
mound the men imitated the movements of the 
bird by crouching low. 

Early in the morning the marraidjirri was 
brought out (see Plate 10). The first dance was the 
wasp dance. While most of the men danced, 
two of their number darted out towards a 
man who held the marraidjirri at arm's length 
on a spear-thrower and pretended that they were 
being attacked at the wasp's nest. The men were 
meant to be looking for yams in the monsoon 
forest and to be driven back by the wasps. The 
women danced at some distance with the common 
rhythmic shuffle known as luku wankain 'ngorro. 
When the dancers reached the jungle fowl nest at 
the end of the path they called out the big names 
associated with the deceased. These were both the 
deceased's maraiin (sacred) names and those of 
the Mildjingi clan. As the calling finished the 
bones were handed over by the deceaseds WB 
(Bulambirri) to the dead man's full sisters. [They 
would not be the custodians of the bones]. 

During the period in which bones are carried 
they are thought to indicate the approach both 
of news bearers and of revenge parties. If the 
bones make a light tap against the bark con- 
tainer this indicates the approach of a person 
with news of some kind. If the tap is loud it 
announces the approach of a war party. 

Once or twice before burial in the hollow 
log coffin, the bones are taken out of the 
tarra and red ochred and this may be associ- 
ated with the change of tarra too. After the 
second red ochring the mother may become 
custodian of the bones. This painting with red 
ochre removes wukundi from the woman who 
carried the bones intially. 

HOLLOW LOG COFFIN 
The holding of the final ceremony is decided 
in this way. The father or his brother, asks the 
relatives carrying the tarra whether they are 
ready to make the coffin. If they agree prepar- 
ations are made for the ceremony. Frequently 
the relatives carrying the tarra feel the need 
to make an excuse to agree and say that they 
are ready because they have carried the bones 
for a long time without help from anybody 
else. The coffin is made by F, EB, YB and 



MEM. NAT. MUS. VICT. 37 PLATE 10 




Dancing with a wasp emblem in a Wullaki buku- 
but (Case 6). 



MEM. NAT. MUS. VICT. 37 PLATE 11 







A hollow log coffin ceremony in Arnhem Bay, 

1937. 



MORTUARY CUSTOMS 



107 



MMB, all members of the deceased's patri- 
moiety. 

Hollow log coffins collectively referred to 
as dupan (hollow) differ in size, ranging from 
1.25 to 4.50 m. and different clans call them 
by different names (see Table 2). 

TABLE 2 

The names used by different groups for their 
hollow log coffins 

CLAN/GROUP NAME 

Ritarango 
Mandjikai Wurrwurr 

Birkilli Djallumbo — associated with a wad- 

ing bird found on the salt pans 

Tjambarapoingo Daimirri — a hollow log thrown by 
an ancestral hero into the sea 
and transformed into a hollow 
stone outside Buckingham Bay 

Liagauwumirri Mululu 

Liagallauwumirri Bardaru — Associated with the crow 
and the milky way 

Marango Kallangurr 

Kanalpingo Larradjadja 

Mildjingi 

Warramirri Kapalla — associated with the fun- 

nel of a steamer and the blow- 
hole of a whale 

Tjapu Larrakit 

However, all are made in the same way. A 
tree which has been hollowed out by termites, 
is cut and cleaned by burning. A circle is in- 
cised 300-600 mm from the top end and 
two small holes cut diagonally opposite each 
other 50-75 mm from the top. The circular in- 
cision is called derong and always painted yel- 
low on yiritja coffins and red on dua coffins. 
The two holes are called eyes and serve to 
commemorate the fact that the coffins w r ere 
originally made by each clan's spirit ancestors 
and in some way personify them. 

After the coffin has been shaped it is moved 
into a large shade where the men work on 
painting it. As the painting nears completion 
the people gather for the final ceremony. Each 
evening there is singing and dancing. On the 
final morning the bones are taken from the 
tarra and covered with red ochre. The skull 
is then painted with the clan design. The coffin 
is brought out and placed at an angle with one 
end supported on a forked stick. Inside a sand 
sculpture of the clan well a close male relative 
of the deceased, often a korrong (FZDS) or 
moralkor (MMBS) breaks the long bones and 



the skull before placing them in the log. Final- 
ly the log is erected (see Plate 11). 

The bones of several people of both moieties 
may be placed in the same coffin. The coffins 
are left standing, eventually decaying and dis- 
appearing without trace. 

References 

Thomson, D. F„ 1961. Marrngitmirri and Kalka— 

medicineman and sorcerer — in Arnhem Land. 

Man 61: 97-102. 
Warner, L., 1958. A black civilization: a social study 

of an Australian tribe. New York: Harper and 

Brothers. 

Notes 

1 The Donald Thomson Ethnographic Collection 
was donated to the University of Melbourne by Mrs 
Thomson following the death of her husband in May 
1970. By agreement the University has lent the Collec- 
tion to the National Museum of Victoria where it is 
now housed. 

The principal purpose in preparing this paper for 
publication is to draw attention to the ethnographic 
riches of the Collection. No compilation of notes 
can do justice to the vision that informed the field- 
work nor create the interpretive synthesis that Thom- 
son had in mind. Inevitably the immense ethnographic 
detail of the fieldnotes can now be appreciated only 
by a series of scholars who will be able to breathe 
interpretive life into different aspects of the many 
but unintegrated details that characterize all fieldnotes. 

This introduction to the Arnhem Land section of 
the Collection has been built on Thomson's notes for 
a paper on 'Kopapoingo Death and Mourning Rituals'. 
Within the general framework of the notes I have 
added descriptions from his fieldnotes as case studies. 
The presentation of both the text and the cases has 
been kept as close to Thomson's own wording as 
possible, but some alteration has been unavoidable 
in the process of converting fieldnotes to continuous 
prose. Further the notes cover several years and 
during that time Thomson's understanding of the 
language and life underwent substantial changes. In 
particular, his spelling of words in the local languages 
altered, so I have standardized on the later forms. A 
number of details in the cases and in particular those 
associated with the wandjur sand sculptures, have had 
to be omitted since the meaning was obscure and 
there was no simple way of setting out the informa- 
tion, some of it possibly deriving significance from its 
location on the page and its position relative to other 
notes. Undoubtedly a scholar with particular know- 
ledge of some of the clans' religious life would be 
able to make sense of some of the notes that have 
been omitted. For this reason any person working 
intensively on a paricular aspect of the mortuary 
customs described here or on the details of symbolism 
in the life of a paricular group will have to consult 
the notes themselves where they will find the odd 
word or phrase that may be of significance to them. 
I have enclosed substantive additions to the text by 
myself in square brackets. 



108 



NICOLAS PETERSON 



I received permission to prepare the paper for 
publication from Mrs Thomson while organizing the 
cataloguing of the ethnographic collection on a grant 
from the Australian Institute of Aboriginal Studies. 
Work on the preparation of this paper has been made 
possible by an appointment as senior Associate in 
Aboriginal and Oceanic Ethnology in the Department 
of History in the Faculty of Arts of the University 
of Melbourne. I am most grateful for the help I 
have received from Mrs Thomson, Margret Darragh, 



Gregory Dening, Alan West and Nancy Williams. 
Special thanks are due to Judith Wiseman for her 
unflagging assistance in all things connected with the 
Collection. 

2 It is uncertain whether this usage of wukundi is 
correct. The primary reference is to places associated 
with death that are tabu in some way (see Case 2). 
Thomson appears to have extended the meaning to 
cover other tabus associated with death. 



*kmat 



SRM 



■ 






8H5 



»?3SE|